Prediction of Hanwoo Cattle Phenotypes from Genotypes Using Machine Learning Methods.

Hanwoo was originally raised for draft purposes, but the increase in local demand for red meat turned that purpose into full-scale meat-type cattle rearing; it is now considered one of the most economically important species and a vital food source for Koreans. The application of genomic selection in Hanwoo breeding programs in recent years was expected to lead to higher genetic progress. However, better statistical methods that can improve the genomic prediction accuracy are required. Hence, this study aimed to compare the predictive performance of three machine learning methods, namely, random forest (RF), extreme gradient boosting method (XGB), and support vector machine (SVM), when predicting the carcass weight (CWT), marbling score (MS), backfat thickness (BFT) and eye muscle area (EMA). Phenotypic and genotypic data (53,866 SNPs) from 7324 commercial Hanwoo cattle that were slaughtered at the age of around 30 months were used. The results showed that the boosting method XGB showed the highest predictive correlation for CWT and MS, followed by GBLUP, SVM, and RF. Meanwhile, the best predictive correlation for BFT and EMA was delivered by GBLUP, followed by SVM, RF, and XGB. Although XGB presented the highest predictive correlations for some traits, we did not find an advantage of XGB or any machine learning methods over GBLUP according to the mean squared error of prediction. Thus, we still recommend the use of GBLUP in the prediction of genomic breeding values for carcass traits in Hanwoo cattle.

Genetic Parameters for Different Measures of Feed Efficiency and Their Relationship to Production Traits in Three Purebred Pigs.

Residual feed intake (RFI) gained attention as a potential alternative to the feed conversion ratio (FCR). Thus, this study aimed to estimate genetic parameters for different feed efficiency (FE) traits (FCR, RFI1 to RFI5) and their genetic correlation to on-test daily weight gain (ADG), backfat (BFT), loin muscle area (LMA), lean percentage (LP), and total feed intake (FI) for 603 Male Duroc (DD), 295 Landrace (LL), and 341 Yorkshire (YY). The common spatial pen effect was also estimated in these traits. Five RFI measures were estimated by regressing daily feed intake on initial testing age (ITA), initial testing weight (IBW), and ADG for RFI1; other models were the same as RFI1 except for additional BFT for RFI2; LMA for RFI3; BFT and LMA for RFI4; BFT, LMA, and average metabolic body weight (AMBW) instead of IBW for RFI5. Genetic parameters estimated using two animal models and the REML method showed moderate heritability for FCR in all breeds (0.22 and 0.28 for DD, 0.31 and 0.39 for LL, 0.17 and 0.22 for YY), low heritability for the majority of RFI measures in DD (0.15 to 0.23) and YY (0.14 to 0.20) and moderate heritability for all RFI measures in LL (0.31 to 0.34). Pen variance explained 7% to 22% for FE and 0% to 9% for production traits' phenotypic variance. The genetic correlation revealed that selection against less complex RFI1 in DD and LL and RFI2 in YY would bring the most advantageous reduction to FI (0.71 for DD, 0.49 for LL, 0.43 YY) without affecting ADG in all breeds (0.06 for DD, -0.11 for LL, 0.05 for YY), decrease in BFT, and increase in LP in DD (0.51 in BFT, -0.77 in LP) and LL (0.45 in BFT, -0.83 in LP). Therefore, inclusion of these breed-specific RFI measures in the future selection criteria would help improve feed efficiency in the swine industry.

Single-Step Genomic Evaluation for Meat Quality Traits, Sensory Characteristics, and Fatty-Acid Composition in Duroc Pigs.

Meat quality and carcass characteristics have gained the attention of breeders due to their increasing economic value. Thus, this study investigated the genomic prediction efficiencies of genomic best linear unbiased prediction (GBLUP) and single-step GBLUP (ssGBLUP) for traits associated with meat quality, sensory characteristics, and fatty-acid composition. A total of 1237 Duroc finishing pigs with 654 individuals genotyped using the Illumina Porcine SNP 60k marker panel were used in this study. Prediction accuracy and bias for GBLUP and ssGBLUP were evaluated using a five-replicates of five-fold cross-validation. Estimation of genetic parameters for traits associated with meat quality, including lightness, yellowness, redness, pH at 24 h post-mortem, moisture content, fat content, water-holding capacity, cooking loss except for shear force (0.19), as well as fatty-acid composition (palmitic, stearic, oleic, linoleic, and linolenic fatty acids), revealed moderate to high heritability estimates ranging from 0.25 to 0.72 and 0.27 to 0.50, respectively, whereas all traits related to sensory characteristics (color, flavor, tenderness, juiciness, and palatability) showed low heritability estimates ranging from 0.08 to 0.14. Meanwhile, assessment of genomic prediction accuracy revealed that ssGBLUP exhibited higher prediction accuracy than GBLUP for meat quality traits, fatty-acid composition, and sensory characteristics, with percentage improvements ranging from 1.90% to 56.07%, 0.73% to 23.21%, and 0.88% to 11.85%, respectively. In terms of prediction bias, ssGBLUP showed less bias estimates than GBLUP for the majority of traits related to meat quality traits, sensory characteristics, and fatty-acid composition of Duroc meat. In this study, ssGBLUP outperformed GBLUP in terms of prediction accuracy and bias for the majority of traits. Through selection and breeding, our findings could be used to promote meat production with improved nutritional value.

Use of gene expression and whole-genome sequence information to improve the accuracy of genomic prediction for carcass traits in Hanwoo cattle.

BACKGROUND: In this study, we assessed the accuracy of genomic prediction for carcass weight (CWT), marbling score (MS), eye muscle area (EMA) and back fat thickness (BFT) in Hanwoo cattle when using genomic best linear unbiased prediction (GBLUP), weighted GBLUP (wGBLUP), and a BayesR model. For these models, we investigated the potential gain from using pre-selected single nucleotide polymorphisms (SNPs) from a genome-wide association study (GWAS) on imputed sequence data and from gene expression information. We used data on 13,717 animals with carcass phenotypes and imputed sequence genotypes that were split in an independent GWAS discovery set of varying size and a remaining set for validation of prediction. Expression data were used from a Hanwoo gene expression experiment based on 45 animals. RESULTS: Using a larger number of animals in the reference set increased the accuracy of genomic prediction whereas a larger independent GWAS discovery dataset improved identification of predictive SNPs. Using pre-selected SNPs from GWAS in GBLUP improved accuracy of prediction by 0.02 for EMA and up to 0.05 for BFT, CWT, and MS, compared to a 50 k standard SNP array that gave accuracies of 0.50, 0.47, 0.58, and 0.47, respectively. Accuracy of prediction of BFT and CWT increased when BayesR was applied with the 50 k SNP array (0.02 and 0.03, respectively) and was further improved by combining the 50 k array with the top-SNPs (0.06 and 0.04, respectively). By contrast, using BayesR resulted in limited improvement for EMA and MS. wGBLUP did not improve accuracy but increased prediction bias. Based on the RNA-seq experiment, we identified informative expression quantitative trait loci, which, when used in GBLUP, improved the accuracy of prediction slightly, i.e. between 0.01 and 0.02. SNPs that were located in genes, the expression of which was associated with differences in trait phenotype, did not contribute to a higher prediction accuracy. CONCLUSIONS: Our results show that, in Hanwoo beef cattle, when SNPs are pre-selected from GWAS on imputed sequence data, the accuracy of prediction improves only slightly whereas the contribution of SNPs that are selected based on gene expression is not significant. The benefit of statistical models to prioritize selected SNPs for estimating genomic breeding values is trait-specific and depends on the genetic architecture of each trait.

Correction: Weighted Genomic Best Linear Unbiased Prediction for Carcass Traits in Hanwoo Cattle. Genes 2019, 10, 1019.

The authors wish to make the following corrections to this paper [...].

Genetic Parameters of Birth Weight and Weaning Weight and Their Relationship with Gestation Length and Age at First Calving in Hanwoo (Bos taurus coreanae).

Hanwoo is one of the most economically important animal species in Korea due to its significant contribution to nutrition. However, the current selection index only focuses to improve carcass traits of Hanwoo. Thus, this study aimed to estimate the genetic parameters of birth weight (BW) and weaning weight (WW) and their genetic and phenotypic relationship to the age at first calving (AFC) and gestation length (GL) of Hanwoo. The genetic parameters for birth weight (BW) and weaning weight (WW) were estimated using the data obtained from 52,173 and 35,800 Hanwoo calves born from February 1998 to March 2017, respectively. Further, these data were used to determine their genetic and phenotypic correlation to age at first calving (AFC) and gestation length (GL). The heritability estimates of BW and WW and correlation coefficients were obtained using the average information restricted maximum likelihood (AIREML) procedure, fit in single and two-trait linear animal models. The estimated direct heritability for BW and WW was moderate (0.22 ± 0.02) and high (0.51 ± 0.03), respectively, while the maternal heritability for both traits was 0.12 ± 0.01 and 0.17 ± 0.01, respectively. The genetic correlation of BW and reproductive traits (AFC and GL) showed a moderate and high positive correlation coefficient of 0.33 ± 0.06 and 0.53 ± 0.02, respectively, while close to zero and low positive phenotypic correlations of 0.06 ± 0.01 and 0.21 ± 0.06 were also observed between the correlated traits, respectively. For the correlation analysis between WW and AFC, both the genetic and phenotypic correlation showed close to zero values of 0.04 ± 0.06 and -0.01 ± 0.01, respectively. Meanwhile, the genetic and phenotypic correlation between WW and GL showed low and negative correlations of -0.09 ± 0.06 and -0.09 ± 0.01, respectively. These obtained estimated variances for BW and WW and their corresponding genetic and phenotypic correlation to AFC and GL can be used as information for genetic improvement and subsequent economic improvement of Hanwoo farming.

RNA Sequencing (RNA-Seq) Based Transcriptome Analysis in Immune Response of Holstein Cattle to Killed Vaccine against Bovine Viral Diarrhea Virus Type I.

Immune response of 107 vaccinated Holstein cattle was initially obtained prior to the ELISA test. Five cattle with high and low bovine viral diarrhea virus (BVDV) type I antibody were identified as the final experimental animals. Blood samples from these animals were then utilized to determine significant differentially expressed genes (DEGs) using the RNA-seq transcriptome analysis and enrichment analysis. Our analysis identified 261 DEGs in cattle identified as experimental animals. Functional enrichment analysis in gene ontology (GO) annotations and Kyoto Encyclopedia of Genes and Genomes (KEGG) pathways revealed the DEGs potentially induced by the inactivated BVDV type I vaccine, and might be responsible for the host immune responses. Our findings suggested that inactivated vaccine induced upregulation of genes involved in different GO annotations, including antigen processing and presentation of peptide antigen (via MHC class I), immune response, and positive regulation of interferon-gamma production. The observed downregulation of other genes involved in immune response might be due to inhibition of toll-like receptors (TLRs) by the upregulation of the Bcl-3 gene. Meanwhile, the result of KEGG pathways revealed that the majority of DEGs were upregulated and enriched to different pathways, including cytokine-cytokine receptor interaction, platelet activation, extracellular matrix (ECM) receptor interaction, hematopoietic cell lineage, and ATP-binding cassette (ABC) transporters. These significant pathways supported our initial findings and are known to play a vital role in shaping adaptive immunity against BVDV type 1. In addition, type 1 diabetes mellitus pathways tended to be significantly enriched. Thus, further studies are needed to investigate the prevalence of type 1 diabetes mellitus in cattle vaccinated with inactivated and live BVDV vaccine.

Genomic Prediction Based on SNP Functional Annotation Using Imputed Whole-Genome Sequence Data in Korean Hanwoo Cattle.

Whole-genome sequence (WGS) data are increasingly being applied into genomic predictions, offering a higher predictive ability by including causal mutations or single-nucleotide polymorphisms (SNPs) putatively in strong linkage disequilibrium with causal mutations affecting the trait. This study aimed to improve the predictive performance of the customized Hanwoo 50 k SNP panel for four carcass traits in commercial Hanwoo population by adding highly predictive variants from sequence data. A total of 16,892 Hanwoo cattle with phenotypes (i.e., backfat thickness, carcass weight, longissimus muscle area, and marbling score), 50 k genotypes, and WGS imputed genotypes were used. We partitioned imputed WGS data according to functional annotation [intergenic (IGR), intron (ITR), regulatory (REG), synonymous (SYN), and non-synonymous (NSY)] to characterize the genomic regions that will deliver higher predictive power for the traits investigated. Animals were assigned into two groups, the discovery set (7324 animals) used for predictive variant detection and the cross-validation set for genomic prediction. Genome-wide association studies were performed by trait to every genomic region and entire WGS data for the pre-selection of variants. Each set of pre-selected SNPs with different density (1000, 3000, 5000, or 10,000) were added to the 50 k genotypes separately and the predictive performance of each set of genotypes was assessed using the genomic best linear unbiased prediction (GBLUP). Results showed that the predictive performance of the customized Hanwoo 50 k SNP panel can be improved by the addition of pre-selected variants from the WGS data, particularly 3000 variants from each trait, which is then sufficient to improve the prediction accuracy for all traits. When 12,000 pre-selected variants (3000 variants from each trait) were added to the 50 k genotypes, the prediction accuracies increased by 9.9, 9.2, 6.4, and 4.7% for backfat thickness, carcass weight, longissimus muscle area, and marbling score compared to the regular 50 k SNP panel, respectively. In terms of prediction bias, regression coefficients for all sets of genotypes in all traits were close to 1, indicating an unbiased prediction. The strategy used to select variants based on functional annotation did not show a clear advantage compared to using whole-genome. Nonetheless, such pre-selected SNPs from the IGR region gave the highest improvement in prediction accuracy among genomic regions and the values were close to those obtained using the WGS data for all traits. We concluded that additional gain in prediction accuracy when using pre-selected variants appears to be trait-dependent, and using WGS data remained more accurate compared to using a specific genomic region.

Weighted Genomic Best Linear Unbiased Prediction for Carcass Traits in Hanwoo Cattle.

The genomic best linear unbiased prediction (GBLUP) method has been widely used in routine genomic evaluation as it assumes a common variance for all single nucleotide polymorphism (SNP). However, this is unlikely in the case of traits influenced by major SNP. Hence, the present study aimed to improve the accuracy of GBLUP by using the weighted GBLUP (WGBLUP), which gives more weight to important markers for various carcass traits of Hanwoo cattle, such as backfat thickness (BFT), carcass weight (CWT), eye muscle area (EMA), and marbling score (MS). Linear and different nonlinearA SNP weighting procedures under WGBLUP were evaluated and compared with unweighted GBLUP and traditional pedigree-based methods (PBLUP). WGBLUP methods were assessed over ten iterations. Phenotypic data from 10,215 animals from different commercial herds that were slaughtered at approximately 30-month-old of age were used. All these animals were genotyped using Illumina Bovine 50k SNP chip and were divided into a training and a validation population by birth date on 1 November 2015. Genomic prediction accuracies obtained in the nonlinearA weighting methods were higher than those of the linear weighting for all traits. Moreover, unlike with linear methods, no sudden drops in the accuracy were noted after the peak was reached in nonlinearA methods. The average accuracies using PBLUP were 0.37, 0.49, 0.40, and 0.37, and 0.62, 0.74, 0.67, and 0.65 using GBLUP for BFT, CWT, EMA, and MS, respectively. Moreover, these accuracies of genomic prediction were further increased to 4.84% and 2.70% for BFT and CWT, respectively by using the nonlinearA method under the WGBLUP model. For EMA and MS, WGBLUP was as accurate as GBLUP. Our results indicate that the WGBLUP using a nonlinearA weighting method provides improved predictions for CWT and BFT, suggesting that the ability of WGBLUP over the other models by weighting selected SNPs appears to be trait-dependent.

Estimation of Genetic Parameters by Single-Trait and Multi-Trait Models for Carcass Traits in Hanwoo Cattle.

Hanwoo breed is preferred in South Korea because of the high standards in marbling and the palatability of its meat. Numerous studies have been conducted and are ongoing to increase the meat production and quality in this beef population. The aim of this study was to estimate and compare genetic parameters for carcass traits using BLUPF90 software. Four models were constructed, single trait pedigree model (STPM), single-trait genomic model (STGM), multi-trait pedigree model (MTPM), and multi-trait genomic model (MTGM), using the pedigree, phenotype, and genomic information of 7991 Hanwoo cattle. Four carcass traits were evaluated: Back fat thickness (BFT), carcass weight (CWT), eye muscle area (EMA), and marbling score (MS). Heritability estimates of 0.40 and 0.41 for BFT, 0.33 and 0.34 for CWT, 0.36 and 0.37 for EMA, and 0.35 and 0.38 for MS were obtained for the single-trait pedigree model and the multi-trait pedigree model, respectively, in Hanwoo. Further, the genomic model showed more improved results compared to the pedigree model, with heritability of 0.39 (CWT), 0.39 (EMA), and 0.46 (MS), except for 0.39 (BFT), which may be due to random events. Utilization of genomic information in the form of single nucleotide polymorphisms (SNPs) has allowed more capturing of the variance from the traits improving the variance components.

Estimation of Genetic Parameters for Reproductive Traits in Hanwoo (Korean Cattle).

Genetic parameters for the reproductive traits of Hanwoo cattle were estimated using data obtained from 15,355 cows in 92 herds across South Korea, which were inseminated from May 1997 to July 2016. An "average information" restricted maximum likelihood (REML) procedure that fit in single-trait and multi-trait animal models was used to estimate the variance components of age at first calving (AFC), calving interval (CI), days open (DO), and gestation length (GL). Results showed the low estimates of heritability for all reproductive traits from both single-trait and multi-trait models. Estimates of heritability for AFC were 0.08 and 0.10 with single-trait and multi-trait models, respectively, while the estimates of heritability using the same animal models ranged from 0.01 to 0.07, 0.01 to 0.09, and 0.10 to 0.16 for CI, DO, and GL, accordingly. While AFC showed positive genetic correlations of 0.52 and 0.46 with CI and DO, respectively, the estimates of genetic and phenotypic correlations of GL with AFC were close to zero. Moreover, phenotypic correlations of GL with CI and DO were also close to zero; however, the corresponding genetic correlations were 0.13 and -0.38 for CI and DO, respectively. These estimated variance components and genetic correlations for reproductive traits can be utilized for genetic improvement programs of Hanwoo cattle.

Genetic parameters and trends for production traits and their relationship with litter traits in Landrace and Yorkshire pigs.

Genetic parameters and trends in the average daily gain (ADG), backfat thickness (BF), loin muscle area (LMA), lean percentage (LP), and age at 90 kg (D90) were estimated for populations of Landrace and Yorkshire pigs. Additionally, the correlations between these production traits and litter traits were estimated. Litter traits included total born (TB) and number born alive (NBA). The data used for this study were obtained from eight farms during 1999 to 2016. Analyses were carried out with a multivariate animal model to estimate genetic parameters for production traits while bivariate analyses were performed to estimate the correlations between production and litter traits. The heritability estimates were 0.52 and 0.43 for ADG; 0.54 and 0.45 for BF; 0.25 and 0.26 for LMA; 0.54 and 0.48 for LP; and 0.56 and 0.46 for D90 in the Landrace and Yorkshire breeds, respectively. The ADG and D90 showed low genetic correlation with BF and LP. The LMA had -0.40, -0.32, 0.49, and 0.39 genetic correlations with ADG, BF, LP, and D90, respectively. Genetic correlations between production and litter traits were generally low, except for the correlations between LMA and TB (-0.23) in Landrace and ADG and TB (-0.16), ADG and NBA (-0.18), D90 and TB (0.19), and D90 and NBA (0.20) in Yorkshire. Genetic trends in production traits were all favorable except for LMA.

Genome wide association study of fatty acid composition in Duroc swine.

OBJECTIVE: Genome wide association study was conducted to identify and validate candidate genes associated with fatty acid composition of pork. METHODS: A total of 480 purebreed Duroc pigs were genotyped using IlluminaPorcine60k bead chips while the association test was implemented following genome-wide rapid association using Mixed Model and Regression-Genomic Control (GRAMMAR-GC) approach. RESULTS: A total of 25, 29, and 16 single nucleotide polymorphisms (SNPs) were significantly associated with stearic (18:0), oleic (18:1) and saturated fatty acids (SFA), respectively. Genome wide significant variants were located on the same region of swine chromosome 14 (SSC14) that spanned from 120 to 124 Mb. Top SNP ALGA008191 was located at 5 kb near the stearoyl-CoA desaturase (SCD) gene. This gene is directly involved in desaturation of stearic acid into oleic acid. General relationship of significant SNPs showed high linkage disequilibrium thus genome-wide signals was attributed to SCD gene. However, understanding the role of other genes like elongation of very long chain fatty acids-3 (ELOVL3) located on this chromosomal segment might help in further understanding of metabolism and biosynthesis of fatty acids. CONCLUSION: Overall, this study provides evidence that validates SCD gene as strong candidate gene associated with fatty acid composition in Duroc pigs. Moreover, this study confirms significant SNPs near ELOVL3 gene.

Variance components estimation for farrowing traits of three purebred pigs in Korea.

OBJECTIVE: This study was conducted to estimate breed-specific variance components for total number born (TNB), number born alive (NBA) and mortality rate from birth through weaning including stillbirths (MORT) of three main swine breeds in Korea. In addition, the importance of including maternal genetic and service sire effects in estimation models was evaluated. METHODS: Records of farrowing traits from 6,412 Duroc, 18,020 Landrace, and 54,254 Yorkshire sows collected from January 2001 to September 2016 from different farms in Korea were used in the analysis. Animal models and the restricted maximum likelihood method were used to estimate variances in animal genetic, permanent environmental, maternal genetic, service sire and residuals. RESULTS: The heritability estimates ranged from 0.072 to 0.102, 0.090 to 0.099, and 0.109 to 0.121 for TNB; 0.087 to 0.110, 0.088 to 0.100, and 0.099 to 0.107 for NBA; and 0.027 to 0.031, 0.050 to 0.053, and 0.073 to 0.081 for MORT in the Duroc, Landrace and Yorkshire breeds, respectively. The proportion of the total variation due to permanent environmental effects, maternal genetic effects, and service sire effects ranged from 0.042 to 0.088, 0.001 to 0.031, and 0.001 to 0.021, respectively. Spearman rank correlations among models ranged from 0.98 to 0.99, demonstrating that the maternal genetic and service sire effects have small effects on the precision of the breeding value. CONCLUSION: Models that include additive genetic and permanent environmental effects are suitable for farrowing traits in Duroc, Landrace, and Yorkshire populations in Korea. This breed-specific variance components estimates for litter traits can be utilized for pig improvement programs in Korea.