Selective genotyping to implement genomic selection in beef cattle breeding.

Genomic selection (GS) plays an essential role in livestock genetic improvement programs. In dairy cattle, the method is already a recognized tool to estimate the breeding values of young animals and reduce generation intervals. Due to the different breeding structures of beef cattle, the implementation of GS is still a challenge and has been adopted to a much lesser extent than dairy cattle. This study aimed to evaluate genotyping strategies in terms of prediction accuracy as the first step in the implementation of GS in beef while some restrictions were assumed for the availability of phenotypic and genomic information. For this purpose, a multi-breed population of beef cattle was simulated by imitating the practical system of beef cattle genetic evaluation. Four genotyping scenarios were compared to traditional pedigree-based evaluation. Results showed an improvement in prediction accuracy, albeit a limited number of animals being genotyped (i.e., 3% of total animals in genetic evaluation). The comparison of genotyping scenarios revealed that selective genotyping should be on animals from both ancestral and younger generations. In addition, as genetic evaluation in practice covers traits that are expressed in either sex, it is recommended that genotyping covers animals from both sexes.

Genomic rotational crossbreeding with advanced optimum contribution selection methods applied to simulated German Angler and German Holstein dairy cattle populations.

Many local dairy cattle breeds are facing genetic extinction due to a large proportion of foreign genes, which have been introgressed in the past. In addition, the performance gap to popular high-yielding breeds is increasing, resulting in a risk of numeric extinction. In the present simulation study, a genomic rotational crossbreeding scheme with the high-yielding German Holstein breed and the numerically small German Angler breed was analysed with the aim to utilize heterosis effects in the crossbred animals. Simultaneously inbreeding was controlled, and the amount of Holstein introgression observed in the Angler breed was reduced. Different scenarios of implementing OCS methods for Angler individuals were evaluated, which differed in their restrictions regarding kinship, native kinship, as well as the amount of genetic contributions from German Holstein. The results showed that rotational crossbreeding can result in superior crossbred offspring compared to the purebred parental lines, whereby OCS methods can simultaneously restrict the increase in inbreeding and keep the Holstein contributions at their current level. However, reducing the amount of migrant contributions while restricting the increase in the native kinship in Angler turned out to be a costly restriction. The reason was that Angler with low genetic contributions from Holsteins tended to have similar Angler ancestors. Consequently, reducing Holstein contributions would considerably increase the native kinship in Angler if it were not constrained. The constraint on the native kinship made a constraint on the conventional kinship superfluous and caused it to increase at a much lower rate than envisaged. This led to both, a high genetic diversity and a low genetic gain. The high genetic diversity in Angler also resulted in lower and oscillating heterosis effects in the crossbred animals. Thus, the reduction of migrant contribution did not increase heterosis effects in the crossbred offspring, and did not result in superior crossbred offspring in general.

Purebred-crossbred genetic parameters for reproductive traits in swine.

For swine breeding programs, testing and selection programs are usually within purebred (PB) populations located in nucleus units that are generally managed differently and tend to have a higher health level than the commercial herds in which the crossbred (CB) descendants of these nucleus animals are expected to perform. This approach assumes that PB animals selected in the nucleus herd will have CB progeny that have superior performance at the commercial level. There is clear evidence that this may not be the case for all traits of economic importance and, thus, including data collected at the commercial herd level may increase the accuracy of selection for commercial CB performance at the nucleus level. The goal for this study was to estimate genetic parameters for five maternal reproductive traits between two PB maternal nucleus populations (Landrace and Yorkshire) and their CB offspring: Total Number Born (TNB), Number Born Alive (NBA), Number Born Alive > 1 kg (NBA > 1 kg), Total Number Weaned (TNW), and Litter Weight at Weaning (LWW). Estimates were based on single-step GBLUP by analyzing any two combinations of a PB and the CB population, and by analyzing all three populations jointly. The genomic relationship matrix between the three populations was generated by using within-population allele frequencies for relationships within a population, and across-population allele frequencies for relationships of the CB with the PB animals. Utilization of metafounders for the two PB populations had no effect on parameter estimates, so the two PB populations were assumed to be genetically unrelated. Joint analysis of two (one PB plus CB) vs. three (both PB and CB) populations did not impact estimates of heritability, additive genetic variance, and genetic correlations. Heritabilities were generally similar between the PB and CB populations, except for LWW and TNW, for which PB populations had about four times larger estimates than CB. Purebred-crossbred genetic correlations (rpc) were larger for Landrace than for Yorkshire, except for NBA > 1 kg. These estimates of rpc indicate that there is potential to improve selection of PB animals for CB performance by including CB information for all traits in the Yorkshire population, but that noticeable additional gains may only occur for NBA > 1 kg and TNW in the Landrace population.

Simultaneous Bayesian estimation of genetic parameters for curves of weight, feed intake, and residual feed intake in beef cattle.

Rates of gain and feed efficiency are important traits in most breeding programs for growing farm animals. The rate of gain (GAIN) is usually expressed over a certain age period and feed efficiency is often expressed as residual feed intake (RFI), defined as observed feed intake (FI) minus expected feed intake based on live weight (WGT) and GAIN. However, the basic traits recorded are always WGT and FI and other traits are derived from these basic records. The aim of this study was to develop a procedure for simultaneous analysis of the basic records and then derive linear traits related to feed efficiency without retorting to any approximation. A bivariate longitudinal random regression model was employed on 13,791 individual longitudinal records of WGT and FI from 2,827 bulls of six different beef breeds tested for their own performance in the period from 7 to 13 mo of age. Genetic and permanent environmental covariance functions for curves of WGT and FI were estimated using Gibbs sampling. Genetic and permanent covariance functions for curves of GAIN were estimated from the first derivative of the function for WGT and finally the covariance functions were extended to curves for RFI, based on the conditional distribution of FI given WGT and GAIN. Furthermore, the covariance functions were extended to include GAIN and RFI defined over different periods of the performance test. These periods included the whole test period as normally used when predicting breeding values for GAIN and RFI for beef bulls. Based on the presented method, breeding values and genetic parameters for derived traits such as GAIN and RFI defined longitudinally or integrated over (parts of) of the test period can be obtained from a joint analysis of the basic records. The resulting covariance functions for WGT, FI, GAIN, and RFI are usually singular but the method presented here does not suffer from the estimation problems associated with defining these traits individually before the genetic analysis. All the results are thus estimated simultaneously, and the set of parameters is consistent.

Genetic parameters and purebred-crossbred genetic correlations for growth, meat quality, and carcass traits in pigs.

Growth, meat quality, and carcass traits are of economic importance in swine breeding. Understanding their genetic basis in purebred (PB) and commercial crossbred (CB) pigs is necessary for a successful breeding program because, although the breeding goal is to improve CB performance, phenotype collection and selection are usually carried out in PB populations housed in biosecure nucleus herds. Thus, the selection is indirect, and the accuracy of selection depends on the genetic correlation between PB and CB performance (rpc). The objectives of this study were to 1) estimate genetic parameters for growth, meat quality, and carcass traits in a PB sire line and related commercial CB pigs and 2) estimate the corresponding genetic correlations between purebred and crossbred performance (rpc). Both objectives were investigated by using pedigree information only (PBLUP) and by combining pedigree and genomic information in a single-step genomic BLUP (ssGBLUP) procedure. Growth rate showed moderate estimates of heritability for both PB and CB based on PBLUP, while estimates were higher in CB based on ssGBLUP. Heritability estimates for meat quality traits were diverse and slightly different based on PB and CB data with both methods. Carcass traits had higher heritability estimates based on PB compared with CB data based on PBLUP and slightly higher estimates for CB data based on ssGBLUP. A wide range of estimates of genetic correlations were obtained among traits within the PB and CB data. In the PB population, estimates of heritabilities and genetic correlations were similar based on PBLUP and ssGBLUP for all traits, while based on the CB data, ssGBLUP resulted in different estimates of genetic parameters with lower SEs. With some exceptions, estimates of rpc were moderate to high. The SE on the rpc estimates was generally large when based on PBLUP due to limited sample size, especially for CBs. In contrast, estimates of rpc based on ssGBLUP were not only more precise but also more consistent among pairs of traits, considering their genetic correlations within the PB and CB data. The wide range of estimates of rpc (less than 0.70 for 7 out of 13 traits) indicates that the use of CB phenotypes recorded on commercial farms, along with genomic information, for selection in the PB population has potential to increase the genetic progress of CB performance.

Effects of Different Strategies for Exploiting Genomic Selection in Perennial Ryegrass Breeding Programs.

Genomic selection (GS) is a potential pathway to accelerate genetic gain for perennial ryegrass (Lolium perenne L.). The main objectives of the present study were to investigate the level of genetic gain and accuracy by applying GS in commercial perennial ryegrass breeding programs. Different scenarios were compared to a conventional breeding program. Simulated scenarios differed in the method of selection and structure of the breeding program. Two scenarios (Phen-Y12 and Phen) for phenotypic selection and three scenarios (GS-Y12, GS and GS-SP) were considered for genomic breeding schemes. All breeding schemes were simulated for 25 cycles. The amount of genetic gain achieved was different across scenarios. Compared to phenotypic scenarios, GS scenarios resulted in substantially larger genetic gain for the simulated traits. This was mainly due to more efficient selection of plots and single plants based on genomic estimated breeding values. Also, GS allows for reduction in waiting time for the availability of the superior genetic materials from previous cycles, which led to at least a doubling or a trebling of genetic gain compared to the traditional program. Reduction in additive genetic variance levels were higher with GS scenarios than with phenotypic selection. The results demonstrated that implementation of GS in ryegrass breeding is possible and presents an opportunity to make very significant improvements in genetic gains.

Genetic variance of metabolomic features and their relationship with body weight and body weight gain in Holstein cattle1.

In recent years, metabolomics has been used to clarify the biology underlying biological samples. In the field of animal breeding, investigating the magnitude of genetic control on the metabolomic profiles of animals and their relationships with quantitative traits adds valuable information to animal improvement schemes. In this study, we analyzed metabolomic features (MFs) extracted from the metabolomic profiles of 843 male Holstein calves. The metabolomic profiles were obtained using nuclear magnetic resonance (NMR) spectroscopy. We investigated 2 alternative methods to control for peak shifts in the NMR spectra, binning and aligning, to determine which approach was the most efficient for assessing genetic variance. Series of univariate analyses were implemented to elucidate the heritability of each MF. Furthermore, records on BW and ADG from 154 to 294 d of age (ADG154-294), 294 to 336 d of age (ADG294-336), and 154 to 336 d of age (ADG154-336) were used in a series of bivariate analyses to establish the genetic and phenotypic correlations with MFs. Bivariate analyses were only performed for MFs that had a heritability significantly different from zero. The heritabilities obtained in the univariate analyses for the MFs in the binned data set were low (<0.2). In contrast, in the aligned data set, we obtained moderate heritability (0.2 to 0.5) for 3.5% of MFs and high heritability (more than 0.5) for 1% of MFs. The bivariate analyses showed that ~12%, ~3%, ~9%, and ~9% of MFs had significant additive genetic correlations with BW, ADG154-294, ADG294-336, and ADG154-336, respectively. In all of the bivariate analyses, the percentage of significant additive genetic correlations was higher than the percentage of significant phenotypic correlations of the corresponding trait. Our results provided insights into the influence of the underlying genetic mechanisms on MFs. Further investigations in this field are needed for better understanding of the genetic relationship among the MFs and quantitative traits.

Balanced selection on purebred and crossbred performance increases gain in crossbreds.

BACKGROUND: Genomic selection can be applied to select purebreds for crossbred performance (CP). The average performance of crossbreds can be considered as the summation of two components, i.e. the breed average (BA) of the parental breeds and heterosis (H) present in crossbreds. Selection of pure breeds for CP based on genomic estimated breeding values for crossbred performance (GEBV-C) or for purebred performance (GEBV-P) may differ in their ability to exploit BA and H and can affect the merit of crossbreds in both the short and long term. Selection based on GEBV-C is beneficial for CP, because H in crossbreds is efficiently exploited, whereas selection on GEBV-P results in more genetic progress in pure breeds, which increases the BA component of CP. To investigate the outcome of selection on GEBV-C and GEBV-P in both the short and long term, a two-way crossbreeding program was simulated to test the following hypotheses: (1) does selection on GEBV-P result in higher long-term CP compared to selection on GEBV-C and (2) does selection on a combination of GEBV-P and GEBV-C lead to more long-term gain in CP than selection on either separately. METHODS: We investigated the performance of crossbreds in a two-way crossbreeding program across 40 generations and considered different criteria to select purebred parents that ranged from selection on purebred performance to selection for CP with different weights on genomic evaluations based on purebred and CP. These criteria were compared under three genetic models to investigate the effects of the amount of dominance variance, absence of over-dominance, and the structure of the reference population on CP, both in the short and long term. RESULTS AND CONCLUSIONS: Although beneficial in the short to medium term, genomic selection in pure breeds on a criterion that specifically targets CP was inferior to selection for purebred performance in the long term. A selection criterion that maximizes a combination of short- and long-term responses in CP, should improve the components that define crossbred merit (i.e., BA and H) simultaneously.

Response to Selection in Finite Locus Models with Nonadditive Effects.

Under the finite-locus model in the absence of mutation, the additive genetic variation is expected to decrease when directional selection is acting on a population, according to quantitative-genetic theory. However, some theoretical studies of selection suggest that the level of additive variance can be sustained or even increased when nonadditive genetic effects are present. We tested the hypothesis that finite-locus models with both additive and nonadditive genetic effects maintain more additive genetic variance (VA) and realize larger medium- to long-term genetic gains than models with only additive effects when the trait under selection is subject to truncation selection. Four genetic models that included additive, dominance, and additive-by-additive epistatic effects were simulated. The simulated genome for individuals consisted of 25 chromosomes, each with a length of 1 M. One hundred bi-allelic QTL, 4 on each chromosome, were considered. In each generation, 100 sires and 100 dams were mated, producing 5 progeny per mating. The population was selected for a single trait (h2 = 0.1) for 100 discrete generations with selection on phenotype or BLUP-EBV. VA decreased with directional truncation selection even in presence of nonadditive genetic effects. Nonadditive effects influenced long-term response to selection and among genetic models additive gene action had highest response to selection. In addition, in all genetic models, BLUP-EBV resulted in a greater fixation of favorable and unfavorable alleles and higher response than phenotypic selection. In conclusion, for the schemes we simulated, the presence of nonadditive genetic effects had little effect in changes of additive variance and VA decreased by directional selection.

Genomic prediction of crossbred performance based on purebred Landrace and Yorkshire data using a dominance model.

BACKGROUND: In pig breeding, selection is usually carried out in purebred populations, although the final goal is to improve crossbred performance. Genomic selection can be used to select purebred parental lines for crossbred performance. Dominance is the likely genetic basis of heterosis and explicitly including dominance in the genomic selection model may be an advantage when selecting purebreds for crossbred performance. Our objectives were two-fold: (1) to compare the predictive ability of genomic prediction models with additive or additive plus dominance effects, when the validation criterion is crossbred performance; and (2) to compare the use of two pure line reference populations to a single combined reference population. METHODS: We used data on litter size in the first parity from two pure pig lines (Landrace and Yorkshire) and their reciprocal crosses. Training was performed (1) separately on pure Landrace (2085) and Yorkshire (2145) sows and (2) the two combined pure lines (4230), which were genotyped for 38 k single nucleotide polymorphisms (SNPs). Prediction accuracy was measured as the correlation between genomic estimated breeding values (GEBV) of pure line boars and mean corrected crossbred-progeny performance, divided by the average accuracy of mean-progeny performance. We evaluated a model with additive effects only (MA) and a model with both additive and dominance effects (MAD). Two types of GEBV were computed: GEBV for purebred performance (GEBV) based on either the MA or MAD models, and GEBV for crossbred performance (GEBV-C) based on the MAD. GEBV-C were calculated based on SNP allele frequencies of genotyped animals in the opposite line. RESULTS: Compared to MA, MAD improved prediction accuracy for both lines. For MAD, GEBV-C improved prediction accuracy compared to GEBV. For Landrace (Yorkshire) boars, prediction accuracies were equal to 0.11 (0.32) for GEBV based on MA, and 0.13 (0.34) and 0.14 (0.36) for GEBV and GEBV-C based on MAD, respectively. Combining animals from both lines into a single reference population yielded higher accuracies than training on each pure line separately. In conclusion, the use of a dominance model increased the accuracy of genomic predictions of crossbred performance based on purebred data.

A crossbred reference population can improve the response to genomic selection for crossbred performance.

BACKGROUND: Breeding goals in a crossbreeding system should be defined at the commercial crossbred level. However, selection is often performed to improve purebred performance. A genomic selection (GS) model that includes dominance effects can be used to select purebreds for crossbred performance. Optimization of the GS model raises the question of whether marker effects should be estimated from data on the pure lines or crossbreds. Therefore, the first objective of this study was to compare response to selection of crossbreds by simulating a two-way crossbreeding program with either a purebred or a crossbred training population. We assumed a trait of interest that was controlled by loci with additive and dominance effects. Animals were selected on estimated breeding values for crossbred performance. There was no genotype by environment interaction. Linkage phase and strength of linkage disequilibrium between quantitative trait loci (QTL) and single nucleotide polymorphisms (SNPs) can differ between breeds, which causes apparent effects of SNPs to be line-dependent. Thus, our second objective was to compare response to GS based on crossbred phenotypes when the line origin of alleles was taken into account or not in the estimation of breeding values. RESULTS: Training on crossbred animals yielded a larger response to selection in crossbred offspring compared to training on both pure lines separately or on both pure lines combined into a single reference population. Response to selection in crossbreds was larger if both phenotypes and genotypes were collected on crossbreds than if phenotypes were only recorded on crossbreds and genotypes on their parents. If both parental lines were distantly related, tracing the line origin of alleles improved genomic prediction, whereas if both parental lines were closely related and the reference population was small, it was better to ignore the line origin of alleles. CONCLUSIONS: Response to selection in crossbreeding programs can be increased by training on crossbred genotypes and phenotypes. Moreover, if the reference population is sufficiently large and both pure lines are not very closely related, tracing the line origin of alleles in crossbreds improves genomic prediction.

Maximizing crossbred performance through purebred genomic selection.

BACKGROUND: In livestock production, many animals are crossbred, with two distinct advantages: heterosis and breed complementarity. Genomic selection (GS) can be used to select purebred parental lines for crossbred performance (CP). Dominance being the likely genetic basis of heterosis, explicitly including dominance in the GS model may be an advantage to select purebreds for CP. Estimated breeding values for CP can be calculated from additive and dominance effects of alleles that are estimated using pure line data. The objective of this simulation study was to investigate the benefits of applying GS to select purebred animals for CP, based on purebred phenotypic and genotypic information. A second objective was to compare the use of two separate pure line reference populations to that of a single reference population that combines both pure lines. These objectives were investigated under two conditions, i.e. either a low or a high correlation of linkage disequilibrium (LD) phase between the pure lines. RESULTS: The results demonstrate that the gain in CP was higher when parental lines were selected for CP, rather than purebred performance, both with a low and a high correlation of LD phase. For a low correlation of LD phase between the pure lines, the use of two separate reference populations yielded a higher gain in CP than use of a single reference population that combines both pure lines. However, for a high correlation of LD phase, marker effects that were estimated using a single combined reference population increased the gain in CP. CONCLUSIONS: Under the hypothesis that performance of crossbred animals differs from that of purebred animals due to dominance, a dominance model can be used for GS of purebred individuals for CP, without using crossbred data. Furthermore, if the correlation of LD phase between pure lines is high, accuracy of selection can be increased by combining the two pure lines into a single reference population to estimate marker effects.

Wool characteristics in the third generation of Arkharmerino × Ghezel and Arkharmerino × Moghani crossbreed sheep.

This investigation was conducted to evaluate the comparative productive performances and effect of some environmental factors on wool characteristics of Arkharmerino × Ghezel (Ar × Gh) and Arkharmerino × Moghani (Ar × Mo) crossbreed sheep. The mid-side fleece samples taken from animals during 2 years (2007-2008) were analyzed. Each sample was measured for average fiber diameter, fiber diameter variability, staple length, proportion of medullated fiber, proportion of kemp, and comfort factor. The comparative values for these fleece characteristics in F(3) generation Ar × Gh were 28.78 ± 0.48 µm, 36.84 ± 1.16%, 11.94 ± 0.35 cm, 7.07 ± 0.93%, 1.02 ± 0.23%, and 68.93%, respectively; while those traits were 29.79 ± 0.43 µm, 41.86 ± 1.16%, 11.96 ± 0.37 cm, 8.13 ± 1.06%, 2.71 ± 0.45%, and 63.33 ± 3.66% for Ar × Mo, respectively. The effects of genotype, sex, birth type, and year of birth were analyzed. Genotype had a significant (P < 0.01) effect on average fiber diameter and proportion of kemp, and Ar × Gh crossbreeds had lower diameter with less proportion of kemp. The two differences in fiber characteristics that were attributable to sex were fiber diameter variability and proportion of medullated fiber, and females had higher measure than males for both traits. Statistical analysis showed that crossbreeding with Arkharmerino generally had positive effects on the fleece favored to be used in the hand woven authentic carpet production.