Predictive performances of animal models using different multibreed relationship matrices in systems with rotational crossbreeding.

BACKGROUND: In livestock breeding, selection for some traits can be improved with direct selection for crossbred performance. However, genetic analyses with phenotypes from crossbred animals require methods for multibreed relationship matrices; especially when some animals are rotationally crossbred. Multiple methods for multibreed relationship matrices exist, but there is a lack of knowledge on how these methods compare for prediction of breeding values with phenotypes from rotationally crossbred animals. Therefore, the objective of this study was to compare models that use different multibreed relationship matrices in terms of ability to predict accurate and unbiased breeding values with phenotypes from two-way rotationally crossbred animals. METHODS: We compared four methods for multibreed relationship matrices: numerator relationship matrices (NRM), García-Cortés and Toro's partial relationship matrices (GT), Strandén and Mäntysaari's approximation to the GT method (SM), and one NRM with metafounders (MF). The methods were compared using simulated data. We simulated two phenotypes; one with and one without dominance effects. Only crossbred animals were phenotyped and only purebred animals were genotyped. RESULTS: The MF and GT methods were the most accurate and least biased methods for prediction of breeding values in rotationally crossbred animals. Without genomic information, all methods were almost equally accurate for prediction of breeding values in purebred animals; however, with genomic information, the MF and GT methods were the most accurate. The GT, MF, and SM methods were the least biased methods for prediction of breeding values in purebred animals. CONCLUSIONS: For prediction of breeding values with phenotypes from rotationally crossbred animals, models using the MF method or the GT method were generally more accurate and less biased than models using the SM method or the NRM method.

Large-scale association study on daily weight gain in pigs reveals overlap of genetic factors for growth in humans.

BACKGROUND: Imputation from genotyping array to whole-genome sequence variants using resequencing of representative reference populations enhances our ability to map genetic factors affecting complex phenotypes in livestock species. The accumulation of knowledge about gene function in human and laboratory animals can provide substantial advantage for genomic research in livestock species. RESULTS: In this study, 201,388 pigs from three commercial Danish breeds genotyped with low to medium (8.5k to 70k) SNP arrays were imputed to whole genome sequence variants using a two-step approach. Both imputation steps achieved high accuracies, and in total this yielded 26,447,434 markers on 18 autosomes. The average estimated imputation accuracy of markers with minor allele frequency ≥ 0.05 was 0.94. To overcome the memory consumption of running genome-wide association study (GWAS) for each breed, we performed within-breed subpopulation GWAS then within-breed meta-analysis for average daily weight gain (ADG), followed by a multi-breed meta-analysis of GWAS summary statistics. We identified 15 quantitative trait loci (QTL). Our post-GWAS analysis strategy to prioritize of candidate genes including information like gene ontology, mammalian phenotype database, differential expression gene analysis of high and low feed efficiency pig and human GWAS catalog for height, obesity, and body mass index, we proposed MRAP2, LEPROT, PMAIP1, ENSSSCG00000036234, BMP2, ELFN1, LIG4 and FAM155A as the candidate genes with biological support for ADG in pigs. CONCLUSION: Our post-GWAS analysis strategy helped to identify candidate genes not just by distance to the lead SNP but also by multiple sources of biological evidence. Besides, the identified QTL overlap with genes which are known for their association with human growth-related traits. The GWAS with this large data set showed the power to map the genetic factors associated with ADG in pigs and have added to our understanding of the genetics of growth across mammalian species.

Single-step genomic evaluation with metafounders for feed conversion ratio and average daily gain in Danish Landrace and Yorkshire pigs.

BACKGROUND: The single-step genomic best linear unbiased prediction (SSGBLUP) method is a popular approach for genetic evaluation with high-density genotype data. To solve the problem that pedigree and genomic relationship matrices refer to different base populations, a single-step genomic method with metafounders (MF-SSGBLUP) was put forward. The aim of this study was to compare the predictive ability and bias of genomic evaluations obtained with MF-SSGBLUP and standard SSGBLUP. We examined feed conversion ratio (FCR) and average daily gain (ADG) in DanBred Landrace (LL) and Yorkshire (YY) pigs using both univariate and bivariate models, as well as the optimal weighting factors (ω), which represent the proportions of the genetic variance not captured by markers, for ADG and FCR in SSGBLUP and MF-SSGBLUP. RESULTS: In general, SSGBLUP and MF-SSGBLUP showed similar predictive abilities and bias of genomic estimated breeding values (GEBV). In the LL population, the predictive ability for ADG reached 0.36 using uni- or bi-variate SSGBLUP or MF-SSGBLUP, while the predictive ability for FCR was highest (0.20) for the bivariate model using MF-SSGBLUP, but differences between analyses were very small. In the YY population, predictive ability for ADG was similar for the four analyses (up to 0.35), while the predictive ability for FCR was highest (0.36) for the uni- and bi-variate MF-SSGBLUP analyses. SSGBLUP and MF-SSGBLUP exhibited nearly the same bias. In general, the bivariate models had lower bias than the univariate models. In the LL population, the optimal ω for ADG was ~ 0.2 in the univariate or bivariate models using SSGBLUP or MF-SSGBLUP, and the optimal ω for FCR was 0.70 and 0.55 for SSGBLUP and MF-SSGBLUP, respectively. In the YY population, the optimal ω ranged from 0.25 to 0. 35 for ADG across the four analyses and from 0.10 to 0.30 for FCR. CONCLUSIONS: Our results indicate that MF-SSGBLUP performed slightly better than SSGBLUP for genomic evaluation. There was little difference in the optimal weighting factors (ω) between SSGBLUP and MF-SSGBLUP. Overall, the bivariate model using MF-SSGBLUP is recommended for single-step genomic evaluation of ADG and FCR in DanBred Landrace and Yorkshire pigs.

Genetic parameters and genomic prediction for feed intake recorded at the group and individual level in different production systems for growing pigs.

BACKGROUND: In breeding programs, recording large-scale feed intake (FI) data routinely at the individual level is costly and difficult compared with other production traits. An alternative approach could be to record FI at the group level since animals such as pigs are normally housed in groups and fed by a shared feeder. However, to date there have been few investigations about the difference between group- and individual-level FI recorded in different environments. We hypothesized that group- and individual-level FI are genetically correlated but different traits. This study, based on the experiment undertaken in purebred DanBred Landrace (L) boars, was set out to estimate the genetic variances and correlations between group- and individual-level FI using a bivariate random regression model, and to examine to what extent prediction accuracy can be improved by adding information of individual-level FI to group-level FI for animals recorded in groups. For both bivariate and univariate models, single-step genomic best linear unbiased prediction (ssGBLUP) and pedigree-based BLUP (PBLUP) were implemented and compared. RESULTS: The variance components from group-level records and from individual-level records were similar. Heritabilities estimated from group-level FI were lower than those from individual-level FI over the test period. The estimated genetic correlations between group- and individual-level FI based on each test day were on average equal to 0.32 (SD = 0.07), and the estimated genetic correlation for the whole test period was equal to 0.23. Our results demonstrate that by adding information from individual-level FI records to group-level FI records, prediction accuracy increased by 0.018 and 0.032 compared with using group-level FI records only (bivariate vs. univariate model) for PBLUP and ssGBLUP, respectively. CONCLUSIONS: Based on the current dataset, our findings support the hypothesis that group- and individual-level FI are different traits. Thus, the differences in FI traits under these two feeding systems need to be taken into consideration in pig breeding programs. Overall, adding information from individual records can improve prediction accuracy for animals with group records.

Prediction of breeding values for group-recorded traits including genomic information and an individually recorded correlated trait.

Records on groups of individuals could be valuable for predicting breeding values when a trait is difficult or costly to measure on single individuals, such as feed intake and egg production. Adding genomic information has shown improvement in the accuracy of genetic evaluation of quantitative traits with individual records. Here, we investigated the value of genomic information for traits with group records. Besides, we investigated the improvement in accuracy of genetic evaluation for group-recorded traits when including information on a correlated trait with individual records. The study was based on a simulated pig population, including three scenarios of group structure and size. The results showed that both the genomic information and a correlated trait increased the accuracy of estimated breeding values (EBVs) for traits with group records. The accuracies of EBV obtained from group records with a size 24 were much lower than those with a size 12. Random assignment of animals to pens led to lower accuracy due to the weaker relationship between individuals within each group. It suggests that group records are valuable for genetic evaluation of a trait that is difficult to record on individuals, and the accuracy of genetic evaluation can be considerably increased using genomic information. Moreover, the genetic evaluation for a trait with group records can be greatly improved using a bivariate model, including correlated traits that are recorded individually. For efficient use of group records in genetic evaluation, relatively small group size and close relationships between individuals within one group are recommended.

Prediction of genetic merit for growth rate in pigs using animal models with indirect genetic effects and genomic information.

BACKGROUND: Several studies have found that the growth rate of a pig is influenced by the genetics of the group members (indirect genetic effects). Accounting for these indirect genetic effects in a selection program may increase genetic progress for growth rate. However, indirect genetic effects are small and difficult to predict accurately. Genomic information may increase the ability to predict indirect genetic effects. Thus, the objective of this study was to test whether including indirect genetic effects in the animal model increases the predictive performance when genetic effects are predicted with genomic relationships. In total, 11,255 pigs were phenotyped for average daily gain between 30 and 94 kg, and 10,995 of these pigs were genotyped. Two relationship matrices were used: a numerator relationship matrix ([Formula: see text]) and a combined pedigree and genomic relationship matrix ([Formula: see text]); and two different animal models were used: an animal model with only direct genetic effects and an animal model with both direct and indirect genetic effects. The predictive performance of the models was defined as the Pearson correlation between corrected phenotypes and predicted genetic levels. The predicted genetic level of a pig was either its direct genetic effect or the sum of its direct genetic effect and the indirect genetic effects of its group members (total genetic effect). RESULTS: The highest predictive performance was achieved when total genetic effects were predicted with genomic information (21.2 vs. 14.7%). In general, the predictive performance was greater for total genetic effects than for direct genetic effects (0.1 to 0.5% greater; not statistically significant). Both types of genetic effects had greater predictive performance when they were predicted with [Formula: see text] rather than [Formula: see text] (5.9 to 6.3%). The difference between predictive performances of total genetic effects and direct genetic effects was smaller when [Formula: see text] was used rather than [Formula: see text]. CONCLUSIONS: This study provides evidence that: (1) corrected phenotypes are better predicted with total genetic effects than with direct genetic effects only; (2) both direct genetic effects and indirect genetic effects are better predicted with [Formula: see text] than [Formula: see text]; (3) using [Formula: see text] rather than [Formula: see text] primarily improves the predictive performance of direct genetic effects.

Genomic diversity revealed by whole-genome sequencing in three Danish commercial pig breeds.

Whole-genome sequencing of 217 animals from three Danish commercial pig breeds (Duroc, Landrace [LL], and Yorkshire [YY]) was performed. Twenty-six million single-nucleotide polymorphisms (SNPs) and 8 million insertions or deletions (indels) were uncovered. Among the SNPs, 493,099 variants were located in coding sequences, and 29,430 were predicted to have a high functional impact such as gain or loss of stop codon. Using the whole-genome sequence dataset as the reference, the imputation accuracy for pigs genotyped with high-density SNP chips was examined. The overall average imputation accuracy for all biallelic variants (SNP and indel) was 0.69, while it was 0.83 for variants with minor allele frequency > 0.1. This study provides whole-genome reference data to impute SNP chip-genotyped animals for further studies to fine map quantitative trait loci as well as improving the prediction accuracy in genomic selection. Signatures of selection were identified both through analyses of fixation and differentiation to reveal selective sweeps that may have had prominent roles during breed development or subsequent divergent selection. However, the fixation indices did not indicate a strong divergence among these three breeds. In LL and YY, the integrated haplotype score identified genomic regions under recent selection. These regions contained genes for olfactory receptors and oxidoreductases. Olfactory receptor genes that might have played a major role in the domestication were previously reported to have been under selection in several species including cattle and swine.

Genetic associations between stayability and longevity in commercial crossbred sows, and stayability in multiplier sows.

Longevity in commercial sows is often selected for through stayability traits measured in purebred animals. However, this may not be justifiable because longevity and stayability may be subject to both genotype by environment interaction (G × E) and genotype by genotype interaction (G × G). This study tested the hypothesis that stayability to service after first parity is more strongly genetically correlated with longevity in commercial herds when stayability is measured in commercial herds rather than multiplier herds. The analysis was based on farrowing- and service-records from 470,824 sows (189,263 multiplier; 281,561 commercial) and 300 herds (156 multiplier; 144 commercial sows). Multiplier sows were either purebred Landrace or Yorkshire and commercial sows were mainly rotationally crossbreds between the two breeds. Commercial longevity was defined as age in days when culled (LongC), and stayability to service after first parity was defined for both commercial sows (StayC) and multiplier sows (StayM). The genetic correlations between LongC, StayC, and StayM were estimated by restricted maximum likelihood using linear mixed models. Genetic parameters were estimated separately for Landrace and Yorkshire. In Landrace, the genetic correlations between LongC and StayC, LongC and StayM, and StayC and StayM were 0.86 ± 0.02, 0.24 ± 0.05, and 0.34 ± 0.06, respectively. In Yorkshire, the genetic correlations between LongC and StayC, LongC and StayM, and StayC and StayM were 0.81 ± 0.03, 0.17 ± 0.05, and 0.18 ± 0.7, respectively. Conclusively, longevity in commercial herds is more strongly correlated with stayability when stayability is measured in commercial herds rather than multiplier herds.

Correction to: A bivariate genomic model with additive, dominance and inbreeding depression effects for sire line and three-way crossbred pigs.

An amendment to this paper has been published and can be accessed via the original article.

A bivariate genomic model with additive, dominance and inbreeding depression effects for sire line and three-way crossbred pigs.

BACKGROUND: Crossbreeding is widely used in pig production because of the benefits of heterosis effects and breed complementarity. Commonly, sire lines are bred for traits such as feed efficiency, growth and meat content, whereas maternal lines are also bred for reproduction and longevity traits, and the resulting three-way crossbred pigs are used for production of meat. The most important genetic basis for heterosis is dominance effects, e.g. removal of inbreeding depression. The aims of this study were to (1) present a modification of a previously developed model with additive, dominance and inbreeding depression genetic effects for analysis of data from a purebred sire line and three-way crossbred pigs; (2) based on this model, present equations for additive genetic variances, additive genetic covariance, and estimated breeding values (EBV) with associated accuracies for purebred and crossbred performances; (3) use the model to analyse four production traits, i.e. ultra-sound recorded backfat thickness (BF), conformation score (CONF), average daily gain (ADG), and feed conversion ratio (FCR), recorded on Danbred Duroc and Danbred Duroc-Landrace-Yorkshire crossbred pigs reared in the same environment; and (4) obtain estimates of genetic parameters, additive genetic correlations between purebred and crossbred performances, and EBV with associated accuracies for purebred and crossbred performances for this data set. RESULTS: Additive genetic correlations (with associated standard errors) between purebred and crossbred performances were equal to 0.96 (0.07), 0.83 (0.16), 0.75 (0.17), and 0.87 (0.18) for BF, CONF, ADG, and FCR, respectively. For BF, ADG, and FCR, the additive genetic variance was smaller for purebred performance than for crossbred performance, but for CONF the reverse was observed. EBV on Duroc boars were more accurate for purebred performance than for crossbred performance for BF, CONF and FCR, but not for ADG. CONCLUSIONS: Methodological developments led to equations for genetic (co)variances and EBV with associated accuracies for purebred and crossbred performances in a three-way crossbreeding system. As illustrated by the data analysis, these equations may be useful for implementation of genomic selection in this system.

Use of Repeated Group Measurements with Drop Out Animals for Variance Component Estimation and Genetic Evaluation: A Simulation Study.

The efficiency of feed utilization plays an important role in animal breeding. However, measuring feed intake (FI) is costly on an individual basis under practical conditions. Using group measurements to model FI could be practically feasible and cost-effective. The objectives of this study were to develop a random regression model based on repeated group measurements with consideration of missing phenotypes caused by drop out animals. Focus is on variance components (VC) estimation and genetic evaluation, and to investigate the effect of group composition on VC estimation and genetic evaluation using simulated datasets. Data were simulated based on individual FI in a pig population. Each individual had measurement on FI at 6 different time points, reflecting 6 different weeks during the test period. The simulated phenotypes consisted of additive genetic, permanent environment, and random residual effects. Additive genetic and permanent environmental effects were both simulated and modeled by first order Legendre polynomials. Three grouping scenarios based on genetic relationships among the group members were investigated: (1) medium within and across pen genetic relationship; (2) high within group relationship; (3) low within group relationship. To investigate the effect of the drop out animals during test period, a proportion (15%) of animals with individual phenotypes was set as the drop out animals, and two drop out scenarios within each grouping scenario were assessed: (1) animals were randomly dropped out; (2) animals with lower phenotypes were dropped out based on the ranking at each time point. The results show that using group measurements yielded similar VCs estimates but with larger SDs compared with the corresponding scenario of using individual measurements. Compared to scenarios without drop out, similar VC estimates were observed when animals were dropped out randomly, whereas reduced VC estimates were observed when animals were dropped out by the ranking of phenotypes. Different grouping scenarios produced similar VC estimates. Compared to scenarios without drop out, there were no loss of accuracies of genetic evaluation for drop out scenarios. However, dropping out animals by the ranking of phenotypes produced larger bias of estimated breeding values compared to the scenario without dropped out animals and scenario of dropping out animals by random. In conclusion, with an optimized group structure, the developed model can properly handle group measurements with drop out animals, and can achieve comparable accuracy of genetic evaluation for traits measured at the group level.

Estimation of variance components and prediction of breeding values based on group records from varying group sizes.

BACKGROUND: Records on groups of individuals rather than on single individuals could be valuable for predicting breeding values (BV) of the traits that are difficult or costly to measure individually, such as feed intake in pigs or beef cattle. Here, we present a model, which handles group records from varying group sizes and involves multiple fixed and random effects, for estimating variance components and predicting BV. Moreover, using simulation, we investigated the efficiency of group records for predicting BV in situations with various group sizes and structures, and factors that affect the trait. RESULTS: The results show that the presented model for group records worked well and that variances estimated from group records with varying group sizes were consistent with those estimated from individual records, but with larger standard errors. Ignoring litter and pen effects had very little or no influence on the accuracy of estimated BV (EBV) obtained from group records. However, ignoring litter effects resulted in biased estimates of additive genetic variance and EBV. The presence of litter and pen effects on phenotypes decreased the accuracy of EBV although the prediction model fitted both effects. Having more littermates in the same pen led to a higher accuracy of EBV. The decay of EBV accuracy with increasing group size was more marked for scenarios with litter and pen effects than without. When litters of six individuals were divided into two pens, accuracies of EBV obtained from group records with a size up to 12 (average 9.6) and up to 24 (average 19.2) were 66.6 and 57.6% of those estimated from individual records in the scenario with litter and pen effects on phenotypes. These percentages reached 77.0 and 68.4% in the scenario without litter and pen effects on phenotypes. CONCLUSIONS: Our results indicate that the model works appropriately for the analysis of group records from varying group sizes. Using group records for genetic evaluation of traits such as feed intake in pig is feasible and the efficiency of the resulting estimates depends on the size and structure of the groups and on the magnitude of the variances for litter and pen effects.

Genomic prediction using models with dominance and imprinting effects for backfat thickness and average daily gain in Danish Duroc pigs.

BACKGROUND: Dominance and imprinting genetic effects have been shown to contribute to genetic variance for certain traits but are usually ignored in genomic prediction of complex traits in livestock. The objectives of this study were to estimate variances of additive, dominance and imprinting genetic effects and to evaluate predictions of genetic merit based on genomic data for average daily gain (DG) and backfat thickness (BF) in Danish Duroc pigs. METHODS: Corrected phenotypes of 8113 genotyped pigs from breeding and multiplier herds were used. Four Bayesian mixture models that differed in the type of genetic effects included: (A) additive genetic effects, (AD) additive and dominance genetic effects, (AI) additive and imprinting genetic effects, and (ADI) additive, dominance and imprinting genetic effects were compared using Bayes factors. The ability of the models to predict genetic merit was compared with regard to prediction reliability and bias. RESULTS: Based on model ADI, narrow-sense heritabilities of 0.18 and 0.31 were estimated for DG and BF, respectively. Dominance and imprinting genetic effects accounted for 4.0 to 4.6 and 1.3 to 1.4 % of phenotypic variance, respectively, which were statistically significant. Across the four models, reliabilities of the predicted total genetic values (GTV, sum of all genetic effects) ranged from 16.1 (AI) to 18.4 % (AD) for DG and from 30.1 (AI) to 31.4 % (ADI) for BF. The least biased predictions of GTV were obtained with model AD, with regression coefficients of corrected phenotypes on GTV equal to 0.824 (DG) and 0.738 (BF). Reliabilities of genomic estimated breeding values (GBV, additive genetic effects) did not differ significantly among models for DG (between 16.5 and 16.7 %); however, for BF, model AD provided a significantly higher reliability (31.3 %) than model A (30.7 %). The least biased predictions of GBV were obtained with model AD with regression coefficients of 0.872 for DG and 0.764 for BF. CONCLUSIONS: Dominance and genomic imprinting effects contribute significantly to the genetic variation of BF and DG in Danish Duroc pigs. Genomic prediction models that include dominance genetic effects can improve accuracy and reduce bias of genomic predictions of genetic merit.

Sparse single-step method for genomic evaluation in pigs.

BACKGROUND: In many animal breeding programs, with the increasing number of genotyped animals, estimation of genomic breeding values by the single-step method is becoming limited by excessive computing requirements. A recently proposed algorithm for proven and young animals (APY) is an approximation that reduces computing time drastically by dividing genotyped animals into core and non-core animals, with only computations for core animals being time-consuming. We hypothesized that choosing core animals based on representing all generations, minimizing the relatedness within the core group, or maximizing the number of genotyped offspring, would result in greater accuracies of estimated breeding values (EBV). METHODS: We compared eight different core groups for the three pig breeds DanAvl Duroc, DanAvl Landrace and DanAvl Yorkshire. These eight sparse approximations of the single-step method were evaluated based on correlations of EBV for genotyped animals obtained from the sparse methods with those obtained from the usual version of the single-step method. We used a single-trait model with daily gain as trait. RESULTS: For core groups that distributed animals across generations, correlations for genotyped animals (from 0.977 to 0.989) were higher than for those that did not distribute core animals across generations (from 0.934 to 0.956). For core groups that maximized the number of genotyped offspring, correlations for genotyped animals (from 0.983 to 0.989) were higher than for other core groups (from 0.934 to 0.981). There was no clear association between low relatedness within the core group and accuracy of approximations. CONCLUSIONS: We found that for core groups that represent all generations and that maximize the number of genotyped offspring, accurate approximations of EBV were obtained. However, we did not find a clear association between accuracy and relatedness within the core group. For the APY method, this is the first study that reports systematic criteria for the creation of core groups that result in more accurate EBV than a similar-sized random core group. Random core groups only ensure across-generation representation. Therefore, we recommend choosing a core group that represents all generations and that maximizes the number of genotyped offspring for single-step genomic evaluation using the APY method.

Increased prediction accuracy using a genomic feature model including prior information on quantitative trait locus regions in purebred Danish Duroc pigs.

BACKGROUND: In animal breeding, genetic variance for complex traits is often estimated using linear mixed models that incorporate information from single nucleotide polymorphism (SNP) markers using a realized genomic relationship matrix. In such models, individual genetic markers are weighted equally and genomic variation is treated as a "black box." This approach is useful for selecting animals with high genetic potential, but it does not generate or utilise knowledge of the biological mechanisms underlying trait variation. Here we propose a linear mixed-model approach that can evaluate the collective effects of sets of SNPs and thereby open the "black box." The described genomic feature best linear unbiased prediction (GFBLUP) model has two components that are defined by genomic features. RESULTS: We analysed data on average daily gain, feed efficiency, and lean meat percentage from 3,085 Duroc boars, along with genotypes from a 60 K SNP chip. In addition information on known quantitative trait loci (QTL) from the animal QTL database was integrated in the GFBLUP as a genomic feature. Our results showed that the most significant QTL categories were indeed biologically meaningful. Additionally, for high heritability traits, prediction accuracy was improved by the incorporation of biological knowledge in prediction models. A simulation study using the real genotypes and simulated phenotypes demonstrated challenges regarding detection of causal variants in low to medium heritability traits. CONCLUSIONS: The GFBLUP model showed increased predictive ability when enough causal variants were included in the genomic feature to explain over 10 % of the genomic variance, and when dilution by non-causal markers was minimal. In the observed data set, predictive ability was increased by the inclusion of prior QTL information obtained outside the training data set, but only for the trait with highest heritability.

Imputation of genotypes in Danish purebred and two-way crossbred pigs using low-density panels.

BACKGROUND: Genotype imputation is commonly used as an initial step in genomic selection since the accuracy of genomic selection does not decline if accurately imputed genotypes are used instead of actual genotypes but for a lower cost. Performance of imputation has rarely been investigated in crossbred animals and, in particular, in pigs. The extent and pattern of linkage disequilibrium differ in crossbred versus purebred animals, which may impact the performance of imputation. In this study, first we compared different scenarios of imputation from 5 K to 8 K single nucleotide polymorphisms (SNPs) in genotyped Danish Landrace and Yorkshire and crossbred Landrace-Yorkshire datasets and, second, we compared imputation from 8 K to 60 K SNPs in genotyped purebred and simulated crossbred datasets. All imputations were done using software Beagle version 3.3.2. Then, we investigated the reasons that could explain the differences observed. RESULTS: Genotype imputation performs as well in crossbred animals as in purebred animals when both parental breeds are included in the reference population. When the size of the reference population is very large, it is not necessary to use a reference population that combines the two breeds to impute the genotypes of purebred animals because a within-breed reference population can provide a very high level of imputation accuracy (correct rate ≥ 0.99, correlation ≥ 0.95). However, to ensure that similar imputation accuracies are obtained for crossbred animals, a reference population that combines both parental purebred animals is required. Imputation accuracies are higher when a larger proportion of haplotypes are shared between the reference population and the validation (imputed) populations. CONCLUSIONS: The results from both real data and pedigree-based simulated data demonstrate that genotype imputation from low-density panels to medium-density panels is highly accurate in both purebred and crossbred pigs. In crossbred pigs, combining the parental purebred animals in the reference population is necessary to obtain high imputation accuracy.

Most of the long-term genetic gain from optimum-contribution selection can be realised with restrictions imposed during optimisation.

BACKGROUND: We tested the hypothesis that optimum-contribution selection (OCS) with restrictions imposed during optimisation realises most of the long-term genetic gain realised by OCS without restrictions. METHODS: We used stochastic simulation to estimate long-term rates of genetic gain realised by breeding schemes that applied OCS without and with restrictions imposed during optimisation, where long-term refers to generations 23 to 25 (approximately). Six restrictions were imposed. Five of these removed solutions from the solution space. The sixth removed records of selection decisions made at earlier selection times. We also simulated a conventional breeding scheme with truncation selection as a reference point. Generations overlapped, selection was for a single trait, and the trait was observed for all selection candidates prior to selection. RESULTS: OCS with restrictions realised 67 to 99% of the additional gain realised by OCS without restrictions, where additional gain was the difference in the long-term rates of genetic gain realised by OCS without restrictions and our reference point with truncation selection. The only exceptions were those restrictions that removed all solutions near the optimum solution from the solution space and the restriction that removed records of selection decisions made at earlier selection times. Imposing these restrictions realised only -12 to 46% of the additional gain. CONCLUSIONS: Most of the long-term genetic gain realised by OCS without restrictions can be realised by OCS with restrictions imposed during optimisation, provided the restrictions do not remove all solutions near the optimum from the solution space and do not remove records of earlier selection decisions. In breeding schemes where OCS cannot be applied optimally because of biological and logistical restrictions, OCS with restrictions provides a useful alternative. Not only does it realise most of the long-term genetic gain, OCS with restrictions enables OCS to be tailored to individual breeding schemes.

Genome-wide association and pathway analysis of feed efficiency in pigs reveal candidate genes and pathways for residual feed intake.

Residual feed intake (RFI) is a complex trait that is economically important for livestock production; however, the genetic and biological mechanisms regulating RFI are largely unknown in pigs. Therefore, the study aimed to identify single nucleotide polymorphisms (SNPs), candidate genes and biological pathways involved in regulating RFI using Genome-wide association (GWA) and pathway analyses. A total of 596 Yorkshire boars with phenotypes for two different measures of RFI (RFI1 and 2) and 60k genotypic data was used. GWA analysis was performed using a univariate mixed model and 12 and 7 SNPs were found to be significantly associated with RFI1 and RFI2, respectively. Several genes such as xin actin-binding repeat-containing protein 2 (XIRP2),tetratricopeptide repeat domain 29 (TTC29),suppressor of glucose, autophagy associated 1 (SOGA1),MAS1,G-protein-coupled receptor (GPCR) kinase 5 (GRK5),prospero-homeobox protein 1 (PROX1),GPCR 155 (GPR155), and FYVE domain containing the 26 (ZFYVE26) were identified as putative candidates for RFI based on their genomic location in the vicinity of these SNPs. Genes located within 50 kbp of SNPs significantly associated with RFI and RFI2 (q-value ≤ 0.2) were subsequently used for pathway analyses. These analyses were performed by assigning genes to biological pathways and then testing the association of individual pathways with RFI using a Fisher's exact test. Metabolic pathway was significantly associated with both RFIs. Other biological pathways regulating phagosome, tight junctions, olfactory transduction, and insulin secretion were significantly associated with both RFI traits when relaxed threshold for cut-off p-value was used (p ≤ 0.05). These results implied porcine RFI is regulated by multiple biological mechanisms, although the metabolic processes might be the most important. Olfactory transduction pathway controlling the perception of feed via smell, insulin pathway controlling food intake might be important pathways for RFI. Furthermore, our study revealed key genes and genetic variants that control feed efficiency that could potentially be useful for genetic selection of more feed efficient pigs.

Genome-wide association and systems genetic analyses of residual feed intake, daily feed consumption, backfat and weight gain in pigs.

BACKGROUND: Feed efficiency is one of the major components determining costs of animal production. Residual feed intake (RFI) is defined as the difference between the observed and the expected feed intake given a certain production. Residual feed intake 1 (RFI1) was calculated based on regression of individual daily feed intake (DFI) on initial test weight and average daily gain. Residual feed intake 2 (RFI2) was as RFI1 except it was also regressed with respect to backfat (BF). It has been shown to be a sensitive and accurate measure for feed efficiency in livestock but knowledge of the genomic regions and mechanisms affecting RFI in pigs is lacking. The study aimed to identify genetic markers and candidate genes for RFI and its component traits as well as pathways associated with RFI in Danish Duroc boars by genome-wide associations and systems genetic analyses. RESULTS: Phenotypic and genotypic records (using the Illumina Porcine SNP60 BeadChip) were available on 1,272 boars. Fifteen and 12 loci were significantly associated (p < 1.52 × 10-6) with RFI1 and RFI2, respectively. Among them, 10 SNPs were significantly associated with both RFI1 and RFI2 implying the existence of common mechanisms controlling the two RFI measures. Significant QTL regions for component traits of RFI (DFI and BF) were detected on pig chromosome (SSC) 1 (for DFI) and 2 for (BF). The SNPs within MAP3K5 and PEX7 on SSC 1, ENSSSCG00000022338 on SSC 9, and DSCAM on SSC 13 might be interesting markers for both RFI measures. Functional annotation of genes in 0.5 Mb size flanking significant SNPs indicated regulation of protein and lipid metabolic process, gap junction, inositol phosphate metabolism and insulin signaling pathway are significant biological processes and pathways for RFI, respectively. CONCLUSIONS: The study detected novel genetic variants and QTLs on SSC 1, 8, 9, 13 and 18 for RFI and indicated significant biological processes and metabolic pathways involved in RFI. The study also detected novel QTLs for component traits of RFI. These results improve our knowledge of the genetic architecture and potential biological pathways underlying RFI; which would be useful for further investigations of key candidate genes for RFI and for development of biomarkers.