Inbreeding in genome-wide selection

Traditional selection methods, such as sib and best linear unbiased prediction (BLUP) selection, which increased genetic gain by increasing accuracy of evaluation have also led to an increased rate of inbreeding per generation (DeltaFG). This is not necessarily the case with genome-wide selection, which also increases genetic gain by increasing accuracy. This paper explains why genome-wide selection reduces DeltaFG when compared with sib and BLUP selection. Genome-wide selection achieves high accuracies of estimated breeding values through better prediction of the Mendelian sampling term component of breeding values. This increases differentiation between sibs and reduces coselection of sibs and DeltaFG. The high accuracy of genome-wide selection is expected to reduce the between family variance and reweigh the emphasis of estimated breeding values of individuals towards the Mendelian sampling term. Moreover, estimation induced intraclass correlations of sibs are expected to be lower in genome-wide selection leading to a further decrease of coselection of sibs when compared with BLUP. Genome-wide prediction of breeding values, therefore, enables increased genetic gain while at the same time reducing DeltaFG when compared with sib and BLUP selection.     

Marker‐assisted selection reduces expected inbreeding but can result in large effects of hitchhiking

We used computer simulations to investigate to what extent true inbreeding, i.e. identity‐by‐descent, is affected by the use of marker‐assisted selection (MAS) relative to traditional best linear unbiased predictions (BLUP) selection. The effect was studied by varying the heritability (h² = 0.04 vs. 0.25), the marker distance (MAS vs. selection on the gene, GAS), the favourable QTL allele effect (α = 0.118 vs. 0.236) and the initial frequency of the favourable QTL allele (p = 0.01 vs. 0.1) in a population resembling the breeding nucleus of a dairy cattle population. The simulated genome consisted of two chromosomes of 100 cM each in addition to a polygenic component. On chromosome 1, a biallelic QTL as well as 4 markers were simulated in linkage disequilibrium. Chromosome 2 was selectively neutral. The results showed that, while reducing pedigree estimated inbreeding, MAS and GAS did not always reduce true inbreeding at the QTL relative to BLUP. MAS and GAS differs from BLUP by increasing the weight on Mendelian sampling terms and thereby lowering inbreeding, while increasing the fixation rate of the favourable QTL allele and thereby increasing inbreeding. The total outcome in terms of inbreeding at the QTL depends on the balance between these two effects. In addition, as a result of hitchhiking, MAS results in extra inbreeding in the region surrounding QTL, which could affect the overall genomic inbreeding.     

Detecting effective starting point of genomic selection by divergent trends from best linear unbiased prediction and single-step genomic best linear unbiased prediction in pigs,beef cattle,and broilers

Genomic selection has been adopted nationally and internationally in different livestock and plant species. However, understanding whether genomic selection has been effective or not is an essential question for both industry and academia. Once genomic evaluation started being used, estimation of breeding values with pedigree best linear unbiased prediction (BLUP) became biased because this method does not consider selection using genomic information. Hence, the effective starting point of genomic selection can be detected in two possible ways including the divergence of genetic trends and Realized Mendelian sampling (RMS) trends obtained with BLUP and single-step genomic BLUP (ssGBLUP). This study aimed to find the start date of genomic selection for a set of economically important traits in three livestock species by comparing trends obtained using BLUP and ssGBLUP. Three datasets were used for this purpose: 1) a pig dataset with 117k genotypes and 1.3M animals in pedigree, 2) an Angus cattle dataset consisted of ~842k genotypes and 11.5M animals in pedigree, and 3) a purebred broiler chicken dataset included ~154k genotypes and 1.3M birds in pedigree were used. The genetic trends for pigs diverged for the genotyped animals born in 2014 for average daily gain (ADG) and backfat (BF). In beef cattle, the trends started diverging in 2009 for weaning weight (WW) and in 2016 for postweaning gain (PWG), with little divergence for birth weight (BTW). In broiler chickens, the genetic trends estimated by ssGBLUP and BLUP diverged at breeding cycle 6 for two out of the three production traits. The RMS trends for the genotyped pigs diverged for animals born in 2014, more for ADG than for BF. In beef cattle, the RMS trends started diverging in 2009 for WW and in 2016 for PWG, with a trivial trend for BTW. In broiler chickens, the RMS trends from ssGBLUP and BLUP diverged strongly for two production traits at breeding cycle 6, with a slight divergence for another trait. Divergence of the genetic trends from ssGBLUP and BLUP indicates the onset of the genomic selection. The presence of trends for RMS indicates selective genotyping, with or without the genomic selection. The onset of genomic selection and genotyping strategies agrees with industry practices across the three species. In summary, the effective start of genomic selection can be detected by the divergence between genetic and RMS trends from BLUP and ssGBLUP.     

Impacts of genotyping strategies on long‐term genetic response in genomic selection

The present study investigated the effects of the choices of animals of reference populations on long‐term responses to genomic selection. Simulated populations comprised 300 individuals and 10 generations of selection practiced for a trait with heritability of 0.1, 0.3 or 0.5. Thirty individuals were randomly selected in the first five generations and selected by estimated breeding values from best linear unbiased prediction (BLUP) and genomic BLUP in the subsequent five generations. The reference populations comprise all animals for all generations (scenario 1), all animals for 6‐10 generations (scenario 2) and 2‐6 generations (scenario 3), and half of the animals for all generations (scenario 4). For all heritability levels, the genetic gains in generation 10 were similar in scenarios 1 and 2. Among scenarios 2 to 4, the highest genetic gains were obtained in scenario 2, with heritabilities of 0.1 and 0.3 as well as scenario 4 with heritability of 0.5. The inbreeding coefficients in scenarios 1, 2 and 4 were lower than those in BLUP, especially within cases with low heritability. These results indicate an appropriate choice of reference population can improve genetic gain and restrict inbreeding even when the reference population size is limited.     

Comparison of genetic improvement for litter size at birth by direct and indirect selection in swine herd

Responses to selection for number of piglets born alive (NBA) by the total number of piglets born (TNB), the NBA, and the NBA plus number of piglets born dead (NBD) were compared using the accuracy of selection and expected genetic gain calculated from the selection index with family information and the real response to selection, using data generated by Monte Carlo computer simulation. The accuracy of selection for NBA selected by TNB was higher than that by NBA only if the genetic correlation between TNB and NBA was close to 1.0, or the value of heritability for the TNB was much larger than that for the NBA. The accuracy of selection for the NBA selected by the combination of the TNB and the NBA was generally highest in the three selection methods in each family structure. Selection by the TNB resulted in the greatest expected genetic gain for the TNB among the selection methods. In the best linear unbiased prediction (BLUP) selection, the genetic gain for the NBA accumulated by the NBA tended to be similar to that accumulated by the combination of the NBA and the NBD, and both genetic gains at generation 10 were significantly larger than that by the TNB (P < 0.001). The accumulated responses selected by the two‐trait animal model BLUP estimated from genetic parameters with errors were similar to those estimated from the true parameters, and there was no significant difference between them. These results indicate that selection by the NBA or by the NBA and the NBD gives more genetic improvement in the NBA than that by the TNB.     

A mating system to reduce inbreeding in selection programmes: Theoretical basis and modification of compensatory mating

Increased rate of inbreeding in selection programmes may have an important effect on mid- and long-term selection response and reproductive performance through reduction in genetic variance and inbreeding depression. Selection on an inherited trait inflates the rate of inbreeding and reduces the effective population size (R obertson 1961; S antiago and C aballero 1995). This can be particularly important in selection based on index with information from relatives (L ush 1947) or best liner unbiased prediction (BLUP) with an animal model (H enderson 1984). In recent years, various methods have been proposed to reduce the rates of inbreeding in selection programmes while keeping genetic gains at the same level. These methods assume various selection and mating strategies. G rundy et al. (1994) showed that the use of biased heritability estimates for BLUP evaluation is one of the simplest and most efficient methods. A direct reduction in the weight on family mean in index selection (T oro and P erez -E nciso 1990), selection for weighted ancestral Mendelian sampling estimates (W oolliams and T hompson 1994; G rundy et al. 1998) and limited use of selected parents (T oro and N ieto 1984; W ei 1995) have also been shown to be efficient methods. Other methods include nonrandom matings of selected parents, such as factorial mating designs (W oolliams 1989), minimum coancestry mating (T oro et al. 1988) and compensatory mating (S antiago and C aballero 1995). Simultaneous optimization of the selection of candidates and their mating allocations has been also considered through mate selection with linear programming techniques (T oro and P erez -E nciso 1990). Among these methods, compensatory mating is a very simple and efficient method (G rundy et al. 1994; S antiago and C aballero 1995; C aballero et al. 1996). This mating system was derived from the theoretical consideration on effective population size under selection (S antiago and C aballero 1995). Although S antiago and C aballero (1995) considered that implementation of this mating could counteract the cumulative effect of selection on the effective population size, the theoretical basis has been little studied. In this paper, the author gives the theoretical basis of compensatory mating. A modification to enhance the effect of compensatory mating is also proposed and the efficiency is examined by stochastic simulation.     

Persistency of accuracy of genomic breeding values for different simulated pig breeding programs in developing countries

Genetic improvement of pigs in tropical developing countries has focused on imported exotic populations which have been subjected to intensive selection with attendant high population‐wide linkage disequilibrium (LD). Presently, indigenous pig population with limited selection and low LD are being considered for improvement. Given that the infrastructure for genetic improvement using the conventional BLUP selection methods are lacking, a genome‐wide selection (GS) program was proposed for developing countries. A simulation study was conducted to evaluate the option of using 60 K SNP panel and observed amount of LD in the exotic and indigenous pig populations. Several scenarios were evaluated including different size and structure of training and validation populations, different selection methods and long‐term accuracy of GS in different population/breeding structures and traits. The training set included previously selected exotic population, unselected indigenous population and their crossbreds. Traits studied included number born alive (NBA), average daily gain (ADG) and back fat thickness (BFT). The ridge regression method was used to train the prediction model. The results showed that accuracies of genomic breeding values (GBVs) in the range of 0.30 (NBA) to 0.86 (BFT) in the validation population are expected if high density marker panels are utilized. The GS method improved accuracy of breeding values better than pedigree‐based approach for traits with low heritability and in young animals with no performance data. Crossbred training population performed better than purebreds when validation was in populations with similar or a different structure as in the training set. Genome‐wide selection holds promise for genetic improvement of pigs in the tropics.     

Use of molecular markers to improve relationship information in the genetic evaluation of beef cattle tick resistance under pedigree‐based models

The selection of genetically superior individuals is conditional upon accurate breeding value predictions which, in turn, are highly depend on how precisely relationship is represented by pedigree. For that purpose, the numerator relationship matrix is essential as a priori information in mixed model equations. The presence of pedigree errors and/or the lack of relationship information affect the genetic gain because it reduces the correlation between the true and estimated breeding values. Thus, this study aimed to evaluate the effects of correcting the pedigree relationships using single‐nucleotide polymorphism (SNP) markers on genetic evaluation accuracies for resistance of beef cattle to ticks. Tick count data from Hereford and Braford cattle breeds were used as phenotype. Genotyping was carried out using a high‐density panel (BovineHD ‐ Illumina^® bead chip with 777 962 SNPs) for sires and the Illumina BovineSNP50 panel (54 609 SNPs) for their progenies. The relationship between the parents and progenies of genotyped animals was evaluated, and mismatches were based on the Mendelian conflicts counts. Variance components and genetic parameters estimates were obtained using a Bayesian approach via Gibbs sampling, and the breeding values were predicted assuming a repeatability model. A total of 460 corrections in relationship definitions were made (Table 1) corresponding to 1018 (9.5%) tick count records. Among these changes, 97.17% (447) were related to the sire's information, and 2.8% (13) were related to the dam's information. We observed 27.2% (236/868) of Mendelian conflicts for sire–progeny genotyped pairs and 14.3% (13/91) for dam–progeny genotyped pairs. We performed 2174 new definitions of half‐siblings according to the correlation coefficient between the coancestry and molecular coancestry matrices. It was observed that higher‐quality genetic relationships did not result in significant differences of variance components estimates; however, they resulted in more accurate breeding values predictions. Using SNPs to assess conflicts between parents and progenies increases certainty in relationships and consequently the accuracy of breeding value predictions of candidate animals for selection. Thus, higher genetic gains are expected when compared to the traditional non‐corrected relationship matrix.     

Use of marker haplotypes to refine covariances among relatives for breeding value estimation

Markers flanking DNA regions, where quantitative trait loci (QTL) have been previously spotted, can be used to trace the common inheritance of major genes for a better definition of covariances among animals. A practical approach to the use of marker data to refine the additive relationship matrix used in the traditional best linear unbiased prediction (BLUP) methodology is presented. The technique allows the number of the mixed model equations to be kept to an animal level, blending polygenic pedigree data with marker haplotype information. The advantage of this marker-assisted selection (MAS) approach over BLUP selection has been assessed through a stochastic simulation. A finite locus model with 32 independent biallelic loci was generated with normally distributed allelic effects. The heritability of the trait, measured on both sexes and on females only, was set to 0.2 and 0.5. Five-allelic markers 2, 10 and 20 cM apart, bracketed the QTL with the largest effect on the trait, accounting for 17% of the genetic variance. The bracketed QTL had two or eight alleles and its position was undefined within the bracket. Results show a moderate 2% advantage of MAS over BLUP in terms of higher genetic response when trait was recorded on both sexes and heritability was 0.2. The benefit is in the short term, but it lasts longer with polyallelic QTL. When the trait was recorded on females only, MAS produced only a small and insignificant genetic gain, but reduced the overall inbreeding in the population. MAS was also inefficient when heritability was 0.5.     

Expected increases in genetic merit from using optimized contributions in two livestock populations of beef cattle and sheep

The expected benefits from optimized selection in real livestock populations were evaluated by applying dynamic selection algorithms to two livestock populations of sheep (Meatlinc) and beef cattle (Aberdeen Angus). In addition, the effects of introducing BLUP evaluations on the population structure, genetic gain, and inbreeding were investigated. The use of BLUP-EBV accelerated the rates of gain in the Meatlinc, but the effects of BLUP evaluations on Aberdeen Angus are not as evident. Although steady increases in the average coefficient of inbreeding (F) were observed, the inbreeding rates (deltaF) before and after the introduction of BLUP evaluations were not significantly different. The observed deltaF in the last generation was 1.0% for Meatlinc and 0.2% for Aberdeen Angus. The application of the dynamic selection algorithms for maximizing genetic gain at a fixed deltaF led to important expected increases in the rate of genetic gain (deltaG). When deltaF was restricted to the value observed in both populations, increments per year in deltaG of 4.6 (i.e., 17%) index units for Meatlinc and 3.5 (i.e., 30%) index units for Aberdeen Angus were found in comparison to the deltaG expected from conventional truncation BLUP selection. More relaxed constraints on deltaF allowed even higher expected increases in deltaG in both populations. This study demonstrates that the optimization tools constitute a potentially highly effective way of managing gain and inbreeding under a broad range of schemes in terms of scale and inbreeding level. No losses in genetic gain were associated with the use of dynamic optimization selection when schemes were compared at the same deltaF.     

Performance of genomic selection under a single-step approach in autochthonous Spanish beef cattle populations

This study evaluated different strategies for implementing a single-step genomic selection programme in two autochthonous Spanish beef cattle populations (Pirenaica—Pi and Rubia Gallega—RG). The strategies were compared in terms of accuracy attained under different scenarios by simulating genomic data over the known genealogy. Several genotyping approaches were tested, as well as, other factors like marker density, effective population size, mutation rate and heritability of the trait. The results obtained showed gains in accuracy with respect to pedigree BLUP evaluation in all cases. The greatest benefit was obtained when the candidates to selection had their genotypes included in the evaluation. Moreover, genotyping the individuals with the most accurate predictions maximized the gains but other suboptimal strategies also yielded satisfactory results. Furthermore, the gains in accuracy increased with the marker density reaching a plateau at around 50,000 markers. Likewise, the effective population size and the mutation rate have also shown an effect, both increasing the accuracy with decreasing values of these population parameters. Finally, the results obtained for the RG population showed greater gains compared to the Pi population, probably attributed to the wider implantation of artificial insemination.     

Investigating pig survival in different production phases using genomic models

Pig survival is an economically important trait with relevant social welfare implications, thus standing out as an important selection criterion for the current pig farming system. We aimed to estimate (co)variance components for survival in different production phases in a crossbred pig population as well as to investigate the benefit of including genomic information through single-step genomic best linear unbiased prediction (ssGBLUP) on the prediction accuracy of survival traits compared with results from traditional BLUP. Individual survival records on, at most, 64,894 crossbred piglets were evaluated under two multi-trait threshold models. The first model included farrowing, lactation, and combined postweaning survival, whereas the second model included nursery and finishing survival. Direct and maternal breeding values were estimated using BLUP and ssGBLUP methods. Furthermore, prediction accuracy, bias, and dispersion were accessed using the linear regression validation method. Direct heritability estimates for survival in all studied phases were low (from 0.02 to 0.08). Survival in preweaning phases (farrowing and lactation) was controlled by the dam and piglet additive genetic effects, although the maternal side was more important. Postweaning phases (nursery, finishing, and the combination of both) showed the same or higher direct heritabilities compared with preweaning phases. The genetic correlations between survival traits within preweaning and postweaning phases were favorable and strong, but correlations between preweaning and postweaning phases were moderate. The prediction accuracy of survival traits was low, although it increased by including genomic information through ssGBLUP compared with the prediction accuracy from BLUP. Direct and maternal breeding values were similarly accurate with BLUP, but direct breeding values benefited more from genomic information. Overall, a slight increase in bias was observed when genomic information was included, whereas dispersion of breeding values was greatly reduced. Combined postweaning survival presented higher direct heritability than in the preweaning phases and the highest prediction accuracy among all evaluated production phases, therefore standing out as a candidate trait for improving survival. Survival is a complex trait with low heritability; however, important genetic gains can still be obtained, especially under a genomic prediction framework.     

Comparison of restricted maximum likelihood and method R for estimating heritability and predicting breeding value under selection

Method R and Restricted Maximum Likelihood (REML) were compared for estimating heritability (h2) and subsequent prediction of breeding values (a) with data subject to selection. A single-trait animal model was used to generate the data and to predict breeding values. The data originated from 10 sires and 100 dams and simulation progressed for 10 overlapping generations. In simulating the data, genetic evaluation used the underlying parameter values and sires and dams were chosen by truncation selection for greatest predicted breeding values. Four alternative pedigree structures were evaluated: complete pedigree information, 50% of phenotypes with sire identities missing, 50% of phenotypes with dam identities missing, and 50% of phenotypes with sire and dams identities missing. Under selection and with complete pedigree data, Method R was a slightly less consistent estimator of h2 than REML. Estimates of h2 by both methods were biased downward when there was selection and loss of pedigree information and were unbiased when no selection was practiced. The empirical mean square error (EMSE) of Method R was several times larger than the EMSE of REML. In a subsequent analysis, different combinations of generations selected and generations sampled were simulated in an effort to disentangle the effects of both factors on Method R estimates of h2. It was observed that Method R overestimated h2 when both the sampling that is intrinsic in the method and the selection occurred in generations 6 to 10. In a final experiment, BLUP(a) were predicted with h2 estimated by either Method R or REML. Subsequently, five more generations of selection were practiced, and the mean square error of prediction (MSEP) of BLUP(a) was calculated with estimated h2 by either method, or the true value of the parameter. The MSEP of empirical BLUP(a) using Method R was greater than the MSEP of empirical BLUP(a) using REML. The latter statistic was closer to prediction error variance of BLUP(a) than the MSEP of empirical BLUP(a) using Method R, indicating that empirical BLUP(a) calculated using REML produced accurate predictions of breeding values under selection. In conclusion, the variability of h2 estimates calculated with Method R was greater than the variability of h2 estimates calculated with REML, with or without selection. Also, the MSEP of EBLUP(a) calculated using estimates of h2 by Method R was larger than MSEP of EBLUP(a) calculated with REML estimates of h2.     

Including an additional systematic environmental effect within a generation in an evaluation model improves accuracy of prediction of breeding values in a closed herd of pigs

The present study evaluated the advantage of mixed‐model techniques over a selection index under different magnitudes of an additional systematic environmental effect (ASEE) in terms of accuracy of prediction and expected genetic gain. The data attempted to simulate a closed herd in a pig breeding program. The base population (G₀) consisted of 10 males and 50 females. Six generations (G₀ to G₅) were selected by using a selection index of three traits without overlapping. Additional systematic environmental constants with four levels in a generation were assigned from a uniform distribution at different ranges. Breeding values of animals in the last generation (G₅) were estimated on the basis of an index of individual phenotype (SI‐U), SI‐U adjusted for ASEE using a least‐squares mean (SI‐A), best linear unbiased prediction using an animal model excluding ASEE (AM‐E), and an animal model including ASEE (AM‐I). Accuracy of prediction and expected genetic gain were larger by the animal model than by the selection index, even if heritability of the traits selected was high and ASEE was set to zero. When ASEE was zero, the accuracy of prediction and expected genetic gain given by SI‐U and AM‐I were similar to those given by SI‐A and AM‐E, respectively. However, the differences in accuracy and expected gain between SI‐U and AI‐A and between AM‐I and AM‐E increased as the range of ASEE increased. It was concluded that selection based on an animal model was more effective than index selection, even if the herd environment was uniform and traits with high heritability were selected, and that it should be always included in an evaluation model, however slight any systematic environmental effect may be in a closed herd.     

Overview of genetic evaluation in dairy cattle

It is costly and time‐consuming to carry out dairy cattle selection on a large experimental scale. For this reason, sire and cow evaluations are almost exclusively based on field data, which are highly affected by a large array of environmental factors. Therefore, it is crucial to adjust for those environmental effects in order to accurately estimate the genetic merits of sires and cows. Index selection is a simple extension of the ordinary least squares under the assumption that the fixed effects are assumed known without error. The mixed‐model equations (MME) of Henderson provide a simpler alternative to the generalized least squares procedure, which is computationally difficult to apply to large data sets. Solution to the MME yields the best linear unbiased estimator of the fixed effects and the best linear unbiased predictor (BLUP) of the random effects. In an animal breeding situation, the random effects such as sire or animal represent the animal's estimated breeding value, which provides a basis for selection decision. The BLUP procedure under sire model assumes random mating between sires and dams. The genetic evaluation procedure has progressed a long way from the dam‐daughter comparison method to animal model, from single trait to multiple trait analysis, and from lactational to test‐day model, to improve accuracy of evaluations. Multiple‐trait evaluation appears desirable because it takes into account the genetic and environmental variance‐covariance of all traits evaluated. For these reasons, multiple‐trait evaluation would reduce bias from selection and achieve a better accuracy of prediction as compared to single‐trait evaluation. The number of traits included in multiple‐trait evaluation should depend upon the breeding goal. Recent advances in molecular and reproductive technologies have created great potential for quantitative geneticists concerning genetic dissection of quantitative traits, and marker‐assisted genetic evaluation and selection.     

The efficiency of genome-wide selection for genetic improvement of net merit

Four methods of selection for net merit comprising 2 correlated traits were compared in this study: 1) EBV-only index (I?), which consists of the EBV of both traits (i.e., traditional 2-trait BLUP selection); 2) GEBV-only index (I?), which comprises the genomic EBV (GEBV) of both traits; 3) GEBV-assisted index (I?), which combines both the EBV and the GEBV of both traits; and 4) GBV-assisted index (I?), which combines both the EBV and the true genomic breeding value (GBV) of both traits. Comparisons of these indices were based on 3 evaluation criteria [selection accuracy, genetic response (ΔH), and relative efficiency] under 64 scenarios that arise from combining 2 levels of genetic correlation (r(G)), 2 ratios of genetic variances between traits, 2 ratios of the genomic variance to total genetic variances for trait 1, 4 accuracies of EBV, and 2 proportions of r(G) explained by the GBV. Both selection accuracy and genetic responses of the indices I?, I?, and I? increased as the accuracy of EBV increased, but the efficiency of the indices I? and I? relative to I? decreased as the accuracy of EBV increased. The relative efficiency of both I? and I? was generally greater when the accuracy of EBV was 0.6 than when it was 0.9, suggesting that the genomic markers are most useful to assist selection when the accuracy of EBV is low. The GBV-assisted index I? was superior to the GEBV-assisted I? in all 64 cases examined, indicating the importance of improving the accuracy of prediction of genomic breeding values. Other parameters being identical, increasing the genetic variance of a high heritability trait would increase the genetic response of the genomic indices (I?, I?, and I?). The genetic responses to I?, I?, and I(4) was greater when the genetic correlation between traits was positive (r(G) = 0.5) than when it was negative (r(G) = -0.5). The results of this study indicate that the effectiveness of the GEBV-assisted index I? is affected by heritability of and genetic correlation between traits, the ratio of genetic variances between traits, the genomic-genetic variance ratio of each index trait, the proportion of genetic correlation accounted for by the genomic markers, and the accuracy of predictions of both EBV and GBV. However, most of these affecting factors are genetic characteristics of a population that is beyond the control of the breeders. The key factor subject to manipulation is to maximize both the proportion of the genetic variance explained by GEBV and the accuracy of both GEBV and EBV. The developed procedures provide means to investigate the efficiency of various genomic indices for any given combination of the genetic factors studied.     

Predicting rates of inbreeding for livestock improvement schemes

This article presents a deterministic method to predict rates of inbreeding (deltaF) for typical livestock improvement schemes. The method is based on a recently developed general theory to predict rates of inbreeding, which uses the concept of long-term genetic contributions. A typical livestock breeding population was modeled, with overlapping generations, BLUP selection, and progeny testing of male selection candidates. Two types of selection were practiced: animals were either selected by truncation on estimated breeding values (EBV) across age classes, or the number of parents selected from each age class was set to a fixed value and truncation selection was practiced within age classes. Bulmer's equilibrium genetic parameters were obtained by iterating on a pseudo-BLUP selection index and deltaF was predicted for the equilibrium situation. Predictions were substantially more accurate than predictions from other available methods, which ignore the effect of selection on deltaF. Predictions were accurate for schemes with up to 20 sires. Predicted deltaF was somewhat too low for schemes with more than 20 sires, which was due to the use of simple linear models to predict genetic contributions. The present method provides a computationally feasible (i.e., deterministic) tool to consider both the rate of inbreeding and the rate of genetic gain when optimizing livestock improvement schemes.     

快速型黄羽肉鸡饲料利用效率性状的基因组选择研究

旨在探究快速型黄羽肉鸡饲料利用效率性状的遗传参数,评估不同方法所得估计育种值的准确性。本研究以自主培育的快速型黄羽肉鸡E系1 923个个体（其中公鸡1 199只,母鸡724只）为研究素材,采用"京芯一号"鸡55K SNP芯片进行基因分型。分别利用传统最佳线性无偏预测（BLUP）、基因组最佳线性无偏预测（GBLUP）和一步法（SSGBLUP）3种方法,基于加性效应模型进行遗传参数估计,通过10倍交叉验证比较3种方法所得估计育种值的准确性。研究性状包括4个生长性状和4个饲料利用效率性状：42日龄体重（BW42D）、56日龄体重（BW56D）、日均增重（ADG）、日均采食量（ADFI）和饲料转化率（FCR）、剩余采食量（RFI）、剩余增长体重（RG）、剩余采食和增长体重（RIG）。结果显示,4个饲料利用效率性状均为低遗传力（0.08~0.20）,其他生长性状为中等偏低遗传力（0.11~0.35）;4个饲料利用效率性状间均为高度遗传相关,RFI、RIG与ADFI间为中度遗传相关,RFI与ADG间无显著相关性,RIG与ADG间为低度遗传相关。本研究在获得SSGBLUP方法的最佳基因型和系谱矩阵权重比基础上,比较8个性状的估计育种值准确性,SSGBLUP方法获得的准确性分别比传统BLUP和GBLUP方法提高3.85%~14.43%和5.21%~17.89%。综上,以RIG为选择指标能够在降低日均采食量的同时保持日均增重,比RFI更适合快速型黄羽肉鸡的选育目标;采用最佳权重比进行SSGBLUP分析,对目标性状估计育种值的预测性能最优,建议作为快速型黄羽肉鸡基因组选择方法。     

Comparison of selection based on phenotype,selection index and best linear unbiased prediction using data from a closed broiler line

1. Selection based on three methods of estimating breeding values, Best Linear Unbiased Prediction (BLUP), selection index (SI), and phenotype (SP) were compared for three traits, juvenile body weight (JW), percentage breast meat yield (BM) and hen‐day rate of egg production (EP) using records provided by a commercial broiler breeding company.

2. Product moment correlations were calculated between breeding values estimated by each method and averaged across sexes. A mean correlation of 0.69 was obtained between selection on SP and BLUP for JW. Mean correlations of 0.88 and 0.68 and 0.87 were obtained between SI and BLUP for the traits JW, EP and BM, respectively.

3. A mean estimated genetic response of 77.7% was obtained with SP for JW relative to BLUP in the absence of restrictions on the selection of close relatives. Estimated genetic responses of 90.7%, 66.9% and 88.4% were obtained by SI relative to BLUP for JW, EP and BM, respectively.

4. Applying restrictions on the selection of close relatives resulted in slight decreases in estimated responses but not in the respective ranking of the selection methods.

5. The results indicate that BLUP could provide commercial breeders with increased selection responses compared to index selection, in particular for traits of low heritability and where relatively few animals possess performance records.     

Efficient selection against categorically scored hip dysplasia in dogs is possible using best linear unbiased prediction and optimum contribution selection: a simulation study

Breeding to reduce the prevalence of categorically scored hip dysplasia (HD), based on phenotypic assessment of radiographic hip status, has had limited success. The aim of this study was to evaluate two selection strategies for improved hip status: truncation selection based on phenotypic record versus best linear unbiased prediction (BLUP), using stochastic simulation and selection scenarios resembling those in real dog populations. In addition, optimum contribution selection (OCS) was evaluated. Two traits were considered: HD (as a categorical trait with five classes and a heritability of 0.45 on the liability scale) and a continuous trait (with a heritability of 0.25) intended to represent other characteristics in the breeding goal. A population structure mimicking that in real dog populations was modelled. The categorical nature of HD caused a considerably lower genetic gain compared to simulating HD as a continuous trait. Genetic gain was larger for BLUP selection than for phenotypic selection in all scenarios. However, BLUP selection resulted in higher rates of inbreeding. By applying OCS, the rate of inbreeding was lowered to about the same level as phenotypic selection but with increased genetic improvement. For efficient selection against HD, use of BLUP breeding values should be prioritized. In small populations, BLUP should be used together with OCS or similar strategy to maintain genetic variation.