Evidence of biases in genetic evaluations due to genomic preselection in dairy cattle

A genomic preselection step of young sires is now often included in dairy cattle breeding schemes. Young sires are selected based on their genomic breeding values. They have better Mendelian sampling contribution so that the assumption of random Mendelian sampling term in genetic evaluations is clearly violated. When these sires and their progeny are evaluated using BLUP, it is feared that estimated breeding values are biased. The effect of genomic selection on genetic evaluations was studied through simulations keeping the structure of the Holstein population in France. The quality of genetic evaluations was assessed by computing bias and accuracy from the difference and correlation between true and estimated breeding values, respectively, and also the mean square error of prediction. Different levels of heritability, selection intensity, and accuracy of genomic evaluation were tested. After only one generation and whatever the scenario, breeding values of preselected young sires and their daughters were significantly underestimated and their accuracy was decreased. Genomic preselection needs to be accounted for in genetic evaluation models.     

Efficiency of multi-breed genomic selection for dairy cattle breeds with different sizes of reference population

Single-breed genomic selection (GS) based on medium single nucleotide polymorphism (SNP) density (~50,000; 50K) is now routinely implemented in several large cattle breeds. However, building large enough reference populations remains a challenge for many medium or small breeds. The high-density BovineHD BeadChip (HD chip; Illumina Inc., San Diego, CA) containing 777,609 SNP developed in 2010 is characterized by short-distance linkage disequilibrium expected to be maintained across breeds. Therefore, combining reference populations can be envisioned. A population of 1,869 influential ancestors from 3 dairy breeds (Holstein, Montbéliarde, and Normande) was genotyped with the HD chip. Using this sample, 50K genotypes were imputed within breed to high-density genotypes, leading to a large HD reference population. This population was used to develop a multi-breed genomic evaluation. The goal of this paper was to investigate the gain of multi-breed genomic evaluation for a small breed. The advantage of using a large breed (Normande in the present study) to mimic a small breed is the large potential validation population to compare alternative genomic selection approaches more reliably. In the Normande breed, 3 training sets were defined with 1,597, 404, and 198 bulls, and a unique validation set included the 394 youngest bulls. For each training set, estimated breeding values (EBV) were computed using pedigree-based BLUP, single-breed BayesC, or multi-breed BayesC for which the reference population was formed by any of the Normande training data sets and 4,989 Holstein and 1,788 Montbéliarde bulls. Phenotypes were standardized by within-breed genetic standard deviation, the proportion of polygenic variance was set to 30%, and the estimated number of SNP with a nonzero effect was about 7,000. The 2 genomic selection (GS) approaches were performed using either the 50K or HD genotypes. The correlations between EBV and observed daughter yield deviations (DYD) were computed for 6 traits and using the different prediction approaches. Compared with pedigree-based BLUP, the average gain in accuracy with GS in small populations was 0.057 for the single-breed and 0.086 for multi-breed approach. This gain was up to 0.193 and 0.209, respectively, with the large reference population. Improvement of EBV prediction due to the multi-breed evaluation was higher for animals not closely related to the reference population. In the case of a breed with a small reference population size, the increase in correlation due to multi-breed GS was 0.141 for bulls without their sire in reference population compared with 0.016 for bulls with their sire in reference population. These results demonstrate that multi-breed GS can contribute to increase genomic evaluation accuracy in small breeds.     

Effect of enlarging the reference population with (un)genotyped animals on the accuracy of genomic selection in dairy cattle

Genomic selection (GS) permits accurate breeding values to be obtained for young animals, shortening the generation interval and accelerating the genetic gain, thereby leading to reduced costs for proven bulls. Genotyping a large number of animals using high-density single nucleotide polymorphism marker arrays is nevertheless expensive, and therefore, a method to reduce the costs of GS is desired. The aim of this study was to investigate an influence of enlarging the reference population, with either genotyped animals or individuals with predicted genotypes, on the accuracy of genomic estimated breeding values. A dairy cattle population was simulated in which proven bulls with 100 daughters were used as a reference population for GS. Phenotypic records were simulated for bulls with heritability equal to the reliability of daughter yield deviations based on 100 daughters. The simulated traits represented heritabilities at the level of individual daughter performance of 0.3, 0.05, and 0.01. Three scenarios were considered in which (1) the reference population consisted of 1,000 genotyped animals, (2) 1,000 ungenotyped animals were added to the reference population, and (3) the 1,000 animals added in scenario 2 were genotyped in addition to the 1,000 animals from scenario 1. Genotypes for ungenotyped animals were predicted with an average accuracy of 0.58. Additionally, an adjustment of the diagonal elements of the G matrix was proposed for animals with predicted genotypes. The accuracy of genomic estimated breeding values for juvenile animals was the highest for the scenario with 2,000 genotyped animals, being 0.90, 0.79, and 0.60 for the heritabilities of 0.3, 0.05, and 0.01, respectively. Accuracies did not differ significantly between the scenario with 1,000 genotyped animals only and the scenario in which 1,000 ungenotyped animals were added and the adjustment of the G matrix was applied. The absence of significant increase in the accuracy of genomic estimated breeding values was attributed to the low accuracy of predicted genotypes. Although the differences were not significant, the difference between scenario 1 and 2 increased with decreasing heritability. Without the adjustment of the diagonal elements of the G matrix, accuracy decreased. Results suggest that inclusion of ungenotyped animals is only expected to enhance the accuracy of GS when the unknown genotypes can be predicted with high accuracy.     

Assessing genomic prediction accuracy for Holstein sires using bootstrap aggregation sampling and leave-one-out cross validation

Since the introduction of genome-enabled prediction for dairy cattle in 2009, genomic selection has markedly changed many aspects of the dairy genetics industry and enhanced the rate of response to selection for most economically important traits. Young dairy bulls are genotyped to obtain their genomic predicted transmitting ability (GPTA) and reliability (REL) values. These GPTA are a main factor in most purchasing, marketing, and culling decisions until bulls reach 5 yr of age and their milk-recorded offspring become available. At that time, daughter yield deviations (DYD) can be compared with the GPTA computed several years earlier. For most bulls, the DYD align well with the initial predictions. However, for some bulls, the difference between DYD and corresponding GPTA is quite large, and published REL are of limited value in identifying such bulls. A method of bootstrap aggregation sampling (bagging) using genomic BLUP (GBLUP) was applied to predict the GPTA of 2,963, 2,963, and 2,803 young Holstein bulls for protein yield, somatic cell score, and daughter pregnancy rate (DPR), respectively. For each trait, 50 bootstrap samples from a reference population comprising 2011 DYD of 8,610, 8,405, and 7,945 older Holstein bulls were used. Leave-one-out cross validation was also performed to assess prediction accuracy when removing specific bulls from the reference population. The main objectives of this study were (1) to assess the extent to which current REL values and alternative measures of variability, such as the bootstrap standard deviation (SD) of predictions, could detect bulls whose daughter performance deviates significantly from early genomic predictions, and (2) to identify factors associated with the reference population that inform about inaccurate genomic predictions. The SD of bootstrap predictions was a mildly useful metric for identifying bulls whose future daughter performance may deviate significantly from early GPTA for protein and DPR. Leave-one-out cross validation allowed us to identify groups of reference population bulls that were influential on other reference population bulls for protein yield and observe their effects on predictions of testing set bulls, as a whole and individually.     

Genomic selection for producer-recorded health event data in US dairy cattle

Emphasizing increased profit through increased dairy cow production has revealed a negative relationship of production with fitness and health traits. Decreased cow health can affect herd profitability through increased rates of involuntary culling and decreased or lost milk sales. The development of genomic selection methodologies, with accompanying substantial gains in reliability for low-heritability traits, may dramatically improve the feasibility of genetic improvement of dairy cow health. Producer-recorded health information may provide a wealth of information for improvement of dairy cow health, thus improving profitability. The principal objective of this study was to use health data collected from on-farm computer systems in the United States to estimate variance components and heritability for health traits commonly experienced by dairy cows. A single-step analysis was conducted to estimate genomic variance components and heritabilities for health events, including cystic ovaries, displaced abomasum, ketosis, lameness, mastitis, metritis, and retained placenta. A blended H matrix was constructed for a threshold model with fixed effects of parity and year-season and random effects of herd-year and sire. The single-step genomic analysis produced heritability estimates that ranged from 0.02 (standard deviation = 0.005) for lameness to 0.36 (standard deviation = 0.08) for retained placenta. Significant genetic correlations were found between lameness and cystic ovaries, displaced abomasum and ketosis, displaced abomasum and metritis, and retained placenta and metritis. Sire reliabilities increased, on average, approximately 30% with the incorporation of genomic data. From the results of these analyses, it was concluded that genetic selection for health traits using producer-recorded data are feasible in the United States, and that the inclusion of genomic data substantially improves reliabilities for these traits.     

Relatedness between numerically small Dutch Red dairy cattle populations and possibilities for multibreed genomic prediction

Red dairy breeds are a valuable cultural and historical asset, and often a source of unique genetic diversity. However, they have difficulties competing with other, more productive, dairy breeds. Improving competitiveness of Red dairy breeds, by accelerating their genetic improvement using genomic selection, may be a promising strategy to secure their long-term future. For many Red dairy breeds, establishing a sufficiently large breed-specific reference population for genomic prediction is often not possible, but may be overcome by adding individuals from another breed. Relatedness between breeds strongly decides the benefit of adding another breed to the reference population. To prioritize among available breeds, the effective number of chromosome segments (M_e) can be used as an indicator of relatedness between individuals from different breeds. The M_e is also an important parameter in determining the accuracy of genomic prediction. The M_e can be estimated both within a population and between 2 populations or breeds, as the reciprocal of the variance of genomic relationships. We investigated relatedness between 6 Dutch Red cattle breeds, Groningen White Headed (GWH), Dutch Friesian (DF), Meuse-Rhine-Yssel (MRY), Dutch Belted (DB), Deep Red (DR), and Improved Red (IR), focusing primarily on the M_e, to predict which of those breeds may benefit from including reference animals of the other breeds. All of these breeds, except MRY, are under high risk of extinction. Our results indicated high variability of M_e, especially between M_e ranging from ～3,500 to ～17,400, indicating different levels of relatedness between the breeds. Two clusters are especially important, one formed by MRY, DR, and IR, and the other comprising DF and DB. Although relatedness between breeds within each of these 2 clusters is high, across-breed genomic prediction is still limited by the current number of genotyped individuals, which for many breeds is low. However, adding MRY individuals would increase the reference population of DR substantially. We estimated that between 11 and 133 individuals from other breeds are needed to achieve accuracy of genomic prediction equivalent to using one additional individual from the same breed. Given the variation in size of the breeds in this study, the benefit of a multibreed reference population is expected to be lower for larger breeds than for the smaller ones.     

Autoregressive single-step test-day model for genomic evaluations of Portuguese Holstein cattle

The multiple-lactation autoregressive test-day (AR) model is the adopted model for the national genetic evaluation of dairy cattle in Portugal. Under this model, animals' permanent environment effects are assumed to follow a first-order autoregressive process over the long (auto-correlations between parities) and short (auto-correlations between test-days within lactation) terms. Given the relevance of genomic prediction in dairy cattle, it is essential to include marker information in national genetic evaluations. In this context, we aimed to evaluate the feasibility of applying the single-step genomic (G)BLUP to analyze milk yield using the AR model in Portuguese Holstein cattle. In total, 11,434,294 test-day records from the first 3 lactations collected between 1994 and 2017 and 1,071 genotyped bulls were used in this study. Rank correlations and differences in reliability among bulls were used to compare the performance of the traditional (A-AR) and single-step (H-AR) models. These 2 modeling approaches were also applied to reduced data sets with records truncated after 2012 (deleting daughters of tested bulls) to evaluate the predictive ability of the H-AR. Validation scenarios were proposed, taking into account young and proven bulls. Average EBV reliabilities, empirical reliabilities, and genetic trends predicted from the complete and reduced data sets were used to validate the genomic evaluation. Average EBV reliabilities for H-AR (A-AR) using the complete data set were 0.52 (0.16) and 0.72 (0.62) for genotyped bulls with no daughters and bulls with 1 to 9 daughters, respectively. These results showed an increase in EBV reliabilities of 0.10 to 0.36 when genomic information was included, corresponding to a reduction of up to 43% in prediction error variance. Considering the 3 validation scenarios, the inclusion of genomic information improved the average EBV reliability in the reduced data set, which ranged, on average, from 0.16 to 0.26, indicating an increase in the predictive ability. Similarly, empirical reliability increased by up to 0.08 between validation tests. The H-AR outperformed A-AR in terms of genetic trends when unproven genotyped bulls were included. The results suggest that the single-step GBLUP AR model is feasible and may be applied to national Portuguese genetic evaluations for milk yield.     

Genomic selection strategies in a small dairy cattle population evaluated for genetic gain and profit

The objective of this study was to evaluate a genomic breeding scheme in a small dairy cattle population that was intermediate in terms of using both young bulls (YB) and progeny-tested bulls (PB). This scheme was compared with a conventional progeny testing program without use of genomic information and, as the extreme case, a juvenile scheme with genomic information, where all bulls were used before progeny information was available. The population structure, cost, and breeding plan parameters were chosen to reflect the Danish Jersey cattle population, being representative for a small dairy cattle population. The population consisted of 68,000 registered cows. Annually, 1,500 bull dams were screened to produce the 500 genotyped bull calves from which 60 YB were selected to be progeny tested. Two unfavorably correlated traits were included in the breeding goal, a production trait (h² = 0.30) and a functional trait (h² = 0.04). An increase in reliability of 5 percentage points for each trait was used in the default genomic scenario. A deterministic approach was used to model the different breeding programs, where the primary evaluation criterion was annual monetary genetic gain (AMGG). Discounted profit was used as an indicator of the economic outcome. We investigated the effect of varying the following parameters: (1) increase in reliability due to genomic information, (2) number of genotyped bull calves, (3) proportion of bull dam sires that are young bulls, and (4) proportion of cow sires that are young bulls. The genomic breeding scheme was both genetically and economically superior to the conventional breeding scheme, even in a small dairy cattle population where genomic information causes a relatively low increase in reliability of breeding values. Assuming low reliabilities of genomic predictions, the optimal breeding scheme according to AMGG was characterized by mixed use of YB and PB as bull sires. Exclusive use of YB for production cows increased AMGG up to 3 percentage points. The results from this study supported our hypothesis that strong interaction effects exist. The strongest interaction effects were obtained between increased reliabilities of genomic estimated breeding values and more intensive use of YB. The juvenile scheme was genetically inferior when the increase in reliability was low (5 percentage points), but became genetically superior at higher reliabilities of genomic estimated breeding values. The juvenile scheme was always superior according to discounted profit because of the shorter generation interval and minimizing costs for housing and feeding waiting bulls.     

The feasibility of using low-density marker panels for genotype imputation and genomic prediction of crossbred dairy cattle of East Africa

Cost-effective high-density (HD) genotypes of livestock species can be obtained by genotyping a proportion of the population using a HD panel and the remainder using a cheaper low-density panel, and then imputing the missing genotypes that are not directly assayed in the low-density panel. The efficacy of genotype imputation can largely be affected by the structure and history of the specific target population and it should be checked before incorporating imputation in routine genotyping practices. Here, we investigated the efficacy of imputation in crossbred dairy cattle populations of East Africa using 4 different commercial single nucleotide polymorphisms (SNP) panels, 3 reference populations, and 3 imputation algorithms. We found that Minimac and a reference population, which included a mixture of crossbred and ancestral purebred animals, provided the highest imputation accuracy compared with other scenarios of imputation. The accuracies of imputation, measured as the correlation between real and imputed genotypes averaged across SNP, were around 0.76 and 0.94 for 7K and 40K SNP, respectively, when imputed up to a 770K panel. We also presented a method to maximize the imputation accuracy of low-density panels, which relies on the pairwise (co)variances between SNP and the minor allele frequency of SNP. The performance of the developed method was tested in a 5-fold cross-validation process where various densities of SNP were selected using the (co)variance method and also by alternative SNP selection methods and then imputed up to the HD panel. The (co)variance method provided the highest imputation accuracies at almost all marker densities, with accuracies being up to 0.19 higher than the random selection of SNP. The accuracies of imputation from 7K and 40K panels selected using the (co)variance method were around 0.80 and 0.94, respectively. The presented method also achieved higher accuracy of genomic prediction at lower densities of selected SNP. The squared correlation between genomic breeding values estimated using imputed genotypes and those from the real 770K HD panel was 0.95 when the accuracy of imputation was 0.64. The presented method for SNP selection is straightforward in its application and can ensure high accuracies in genotype imputation of crossbred dairy populations in East Africa.     

Genomic prediction of crossbred dairy cattle in Tanzania: A route to productivity gains in smallholder dairy systems

Selection based on genomic predictions has become the method of choice for genetic improvement in dairy cattle. This offers huge opportunity for developing countries with little or no pedigree data, and preliminary studies have shown promising results. The African Dairy Genetic Gains (ADGG) project initiated a digital system of dairy performance data collection, accompanied by genotyping in Tanzania in 2016. Currently, ADGG has the largest body of dairy performance data generated in East Africa from a smallholder dairy system. This study examines the use of genomic best linear unbiased prediction (GBLUP) and single-step (ss)GBLUP for the estimation of genetic parameters and accuracy of genomic prediction for daily milk yield and body weight in Tanzania. The estimates of heritability for daily milk yield from GBLUP and ssGBLUP were essentially the same, at 0.12 ± 0.03. The heritability estimates for daily milk yield averaged over the whole lactation from random regression model (RRM) GBLUP or ssGBLUP were 0.22 and 0.24, respectively. The heritability of body weight from GBLUP was 0.24 ± 04 but was 0.22 ± 04 from the ssGBLUP analysis. Accuracy of genomic prediction for milk yield from a forward validation was 0.57 for GBLUP based on fixed regression model or 0.55 from an RRM. Corresponding estimates from ssGBLUP were 0.59 and 0.53, respectively. Accuracy for body weight, however, was much higher at 0.83 from GBLUP and 0.77 for ssGBLUP. The moderate to high levels of accuracy of genomic prediction (0.53–0.83) obtained for milk yield and body weight indicate that selection on the basis of genomic prediction is feasible in smallholder dairy systems and most probably the only initial possible pathway to implementing sustained genetic improvement programs in such systems.     

Inclusion of herdmate data improves genomic prediction for milk-production and feed-efficiency traits within North American dairy herds

Genomic data are widely available in the dairy industry and provide a cost-effective means of predicting genetic merit to inform selection decisions and increase genetic gains. As more dairy farms adopt genomic selection practices, dairy producers will soon have genomic data available on all of the animals within their herds. This is a very rich, but currently underused, source of information. Herdmates provide an excellent indication of how a selection candidate's genetics will perform within a given herd, noting that herdmates often include close relatives that share a similar environment. The study objective was to evaluate the utility of incorporating herdmate data into genomic predictions in a data set composed of 3,303 Holsteins from one herd in Canada and 6 herds throughout the United States. Within-herd prediction accuracy was assessed for milk-production and feed-efficiency traits determined from genomic best linear unbiased prediction under 4 different scenarios. Scenario 1 did not include herdmates in the training population. Scenarios 2 through 4 included herdmates in the training population, and scenarios 3 and 4 also included modeling of herd-specific marker effects. Leave-one-out cross validation was used to maximize the number of herdmates in the training population in scenarios 2 through 4, while maintaining constant training population size with scenario 1. Results from the present study reveal the importance of incorporating herdmate data into genomic evaluations. Inclusion of herdmates in the training population improved mean within-herd prediction accuracy for milk-production traits (± standard error) by 0.08 ± 0.03 (milk yield), 0.07 ± 0.03 (fat percentage), and 0.05 ± 0.01 (protein percentage) and feed-efficiency traits by 0.07 ± 0.02 (milk energy), 0.03 ± 0.02 (DMI), and 0.08 ± 0.01 (metabolic body weight). Modeling herd-specific marker effects further improved mean within-herd prediction accuracy for milk yield and energy by 0.03 ± 0.01 and 0.02 ± 0.01, respectively. Herds with higher within-herd heritability and low genomic correlation with the remaining herds benefitted most from the inclusion of herdmate data.     

Comparison of genomic predictions for lowly heritable traits using multi-step and single-step genomic best linear unbiased predictor in Holstein cattle

The success and sustainability of a breeding program incorporating genomic information is largely dependent on the accuracy of predictions. For low heritability traits, large training populations are required to achieve high accuracies of genomic estimated breeding values (GEBV). By including genotyped and nongenotyped animals simultaneously in the evaluation, the single-step genomic BLUP (ssGBLUP) approach has the potential to deliver more accurate and less biased genomic evaluations. The aim of this study was to compare the accuracy and bias of genomic predictions for various traits in Canadian Holstein cattle using ssGBLUP and multi-step genomic BLUP (msGBLUP) under different strategies, such as (1) adding genomic information of cows in the analysis, (2) testing different adjustments of the genomic relationship matrix, and (3) using a blending approach to obtain GEBV from msGBLUP. The following genomic predictions were evaluated regarding accuracy and bias: (1) GEBV estimated by ssGBLUP; (2) direct genomic value estimated by msGBLUP with polygenic effects of 5 and 20%; and (3) GEBV calculated by a blending approach of direct genomic value with estimated breeding values using polygenic effects of 5 and 20%. The effect of adding genomic information of cows in the evaluation was also assessed for each approach. When genomic information was included in the analyses, the average improvement in observed reliability of predictions was observed to be 7 and 13 percentage points for reproductive and workability traits, respectively, compared with traditional BLUP. Absolute deviation from 1 of the regression coefficient of the linear regression of de-regressed estimated breeding values on genomic predictions went from 0.19 when using traditional BLUP to 0.22 when using the msGBLUP method, and to 0.14 when using the ssGBLUP method. The use of polygenic weight of 20% in the msGBLUP slightly improved the reliability of predictions, while reducing the bias. A similar trend was observed when a blending approach was used. Adding genomic information of cows increased reliabilities, while decreasing bias of genomic predictions when using the ssGBLUP method. Differences between using a training population with cows and bulls or with only bulls for the msGBLUP method were small, likely due to the small number of cows included in the analysis. Predictions for lowly heritable traits benefit greatly from genomic information, especially when all phenotypes, pedigrees, and genotypes are used in a single-step approach.     

Evaluating the performance of machine learning methods and variable selection methods for predicting difficult-to-measure traits in Holstein dairy cattle using milk infrared spectral data

Fourier-transform infrared (FTIR) spectroscopy is a powerful high-throughput phenotyping tool for predicting traits that are expensive and difficult to measure in dairy cattle. Calibration equations are often developed using standard methods, such as partial least squares (PLS) regression. Methods that employ penalization, rank-reduction, and variable selection, as well as being able to model the nonlinear relations between phenotype and FTIR, might offer improvements in predictive ability and model robustness. This study aimed to compare the predictive ability of 2 machine learning methods, namely random forest (RF) and gradient boosting machine (GBM), and penalized regression against PLS regression for predicting 3 phenotypes differing in terms of biological meaning and relationships with milk composition (i.e., phenotypes measurable directly and not directly in milk, reflecting different biological processes which can be captured using milk spectra) in Holstein-Friesian cattle under 2 cross-validation scenarios. The data set comprised phenotypic information from 471 Holstein-Friesian cows, and 3 target phenotypes were evaluated: (1) body condition score (BCS), (2) blood β-hydroxybutyrate (BHB, mmol/L), and (3) κ-casein expressed as a percentage of nitrogen (κ-CN, % N). The data set was split considering 2 cross-validation scenarios: samples-out random in which the population was randomly split into 10-folds (8-folds for training and 1-fold for validation and testing); and herd/date-out in which the population was randomly assigned to training (70% herd), validation (10%), and testing (20% herd) based on the herd and date in which the samples were collected. The random grid search was performed using the training subset for the hyperparameter optimization and the validation set was used for the generalization of prediction error. The trained model was then used to assess the final prediction in the testing subset. The grid search for penalized regression evidenced that the elastic net (EN) was the best regularization with increase in predictive ability of 5%. The performance of PLS (standard model) was compared against 2 machine learning techniques and penalized regression using 2 cross-validation scenarios. Machine learning methods showed a greater predictive ability for BCS (0.63 for GBM and 0.61 for RF), BHB (0.80 for GBM and 0.79 for RF), and κ-CN (0.81 for GBM and 0.80 for RF) in samples-out cross-validation. Considering a herd/date-out cross-validation these values were 0.58 (GBM and RF) for BCS, 0.73 (GBM and RF) for BHB, and 0.77 (GBM and RF) for κ-CN. The GBM model tended to outperform other methods in predictive ability around 4%, 1%, and 7% for EN, RF, and PLS, respectively. The prediction accuracies of the GBM and RF models were similar, and differed statistically from the PLS model in samples-out random cross-validation. Although, machine learning techniques outperformed PLS in herd/date-out cross-validation, no significant differences were observed in terms of predictive ability due to the large standard deviation observed for predictions. Overall, GBM achieved the highest accuracy of FTIR-based prediction of the different phenotypic traits across the cross-validation scenarios. These results indicate that GBM is a promising method for obtaining more accurate FTIR-based predictions for different phenotypes in dairy cattle.     

Effect of synchronized breeding on genetic evaluations of fertility traits in dairy cattle

Estrus detection has become more difficult over the years due to decreases in the estrus expression of high-producing dairy cows, and increased herd sizes and animal density. Through the use of hormonal synchronization protocols, also known as timed artificial insemination (TAI) protocols, it is possible to alleviate some of the challenges associated with estrus detection. However, TAI masks cows' fertility performance, resulting in an unfair comparison of treated animals and innately fertile animals. Consequently, genetically inferior and superior cows show similar phenotypes, making it difficult to distinguish between them. As genetic programs rely on the collection of accurate phenotypic data, phenotypes collected on treated animals likely add bias to genetic evaluations. In this study, to assess the effect of TAI, the rank correlation of bulls for a given trait using only TAI records were compared with the same trait using only heat detection records. A total of 270,434 records from 192,539 animals split across heifers, first and second parity cows were analyzed for the traits: calving to first service, first service to conception, and days open. Results showed large reranking across all traits and parities between bulls compared based on either having only TAI records or only heat detection records, suggesting that a bias does indeed exist. Large reranking was also observed for both the heat detection and TAI groups among the top 100 bulls in the control group, which included all records. Furthermore, breeding method was added to the model to assess its effect on bull ranking. However, there were only minor changes in the rank correlations between scenario groups. Therefore, more complex methods to account for the apparent bias created by TAI should be investigated; for this, the method by which these data are collected needs to be improved through creating a standardized way of recording breeding codes. Though the results of this study suggest the presence of bias within current fertility evaluations, additional research is required to confirm the findings of this study, including looking at high-reliability bulls specifically, to determine if the levels of reranking remain. Future studies should also aim to understand the potential genetic differences between the fertility traits split via management technology, possibly in a multiple-trait analysis.     

Genotyping strategies for maximizing genomic information in evaluations of the Latxa dairy sheep breed

Genomic selection has been implemented over the years in several livestock species, due to the achievable higher genetic progress. The use of genomic information in evaluations provides better prediction accuracy than do pedigree-based evaluations, and the makeup of the genotyped population is a decisive point. The aim of this work is to compare the effect of different genotyping strategies (number and type of animals) on the prediction accuracy for dairy sheep Latxa breeds. A simulation study was designed based on the real data structure of each population, and the phenotypic and genotypic data obtained were used in genetic (BLUP) and genomic (single-step genomic BLUP) evaluations of different genotyping strategies. The genotyping of males was beneficial when they were genetically connected individuals and if they had daughters with phenotypic records. Genotyping females with their own lactation records increased prediction accuracy, and the connection level has less relevance. The differences in genotyping females were independent of their estimated breeding value. The combined genotyping of males and females provided intermediate accuracy results regardless of the female selection strategy. Therefore, assuming that genotyping rams is interesting, the incorporation of genotyped females would be beneficial and worthwhile. The benefits of genotyping individuals from various generations were highlighted, although it was also possible to gain prediction accuracy when historic individuals were not considered. Greater genotyped population sizes resulted in more accuracy, even if the increase seems to reach a plateau.     

Improving accuracy of bulls' predicted genomic breeding values for fertility using daughters' milk progesterone profiles

The main objective of this study was to investigate the benefit of accuracy of genomic prediction when combining records for an intermediate physiological phenotype in a training population with records for a traditional phenotype. Fertility was used as a case study, where commencement of luteal activity (C-LA) was the physiological phenotype, whereas the interval from calving to first service and calving interval were the traditional phenotypes. The potential accuracy of across-country genomic prediction and optimal recording strategies of C-LA were also investigated in terms of the number of farms and number of repeated records for C-LA. Predicted accuracy was obtained by estimating population parameters for the traits in a data set of 3,136 Holstein Friesian cows with 8,080 lactations and using a deterministic prediction equation. The effect of genetic correlation, heritability, and reliability of C-LA on the accuracy of genomic prediction were investigated. When the existing training population was 10,000 bulls with reliable estimated breeding value for the traditional trait, predicted accuracy for the physiological trait increased from 0.22 to 0.57 when 15,000 cows with C-LA records were added to the bull training population; but, when the interest was in predicting the traditional trait, we found no benefit from the additional recording. When the genetic correlation was higher between the physiological and traditional traits (0.7 instead of 0.3), accuracy increased less when adding the 15.000 cows with C-LA (from 0.51 to 0.63). In across-country predictions, we observed little to no increase in accuracy of the intermediate physiological phenotype when the training population from Sweden was large, but when accuracy increased the training population was small (200 cows), from 0.19 to 0.31 when 15,000 cows were added from the Netherlands (genetic correlation of 0.5 between countries), and from 0.19 to 0.48 for genetic correlation of 0.9. The predicted accuracy initially increased substantially when recording on the same farm was extended and multiple C-LA records per cow were used in prediction compared with single records; that is, accuracy increased from 0.33 with single records to 0.38 with multiple records (on average 1.6 records per cow) from 2 yr of recording C-LA. But, when the number C-LA per cow increased beyond 2 yr of recording, we noted no substantial benefit in accuracy from multiple records. For example, for 5 yr of recording (on average 2.5 records per cow), accuracy was 0.47; on doubling the recording period to 10 yr (on average 3.1 records per cow), accuracy increased by 0.07 units, whereas when C-LA was recorded for 15 yr (on average 3.3 records per cow) accuracy increased only by 0.05 units. Therefore, for genomic prediction using expensive equipment to record traits for training populations, it is important to optimize the recording strategy. The focus should be on recording more cows rather than continuous recording on the same cows.     

Identifying genetic variants and pathways influencing daughter averages for twinning in North American Holstein cattle and evaluating the potential for genomic selection

Multiple birth in dairy cattle is a detrimental trait both economically for producers and for animal health. Genetics of twinning is complex and has led to several quantitative trait loci regions being associated with increased twinning. To identify variants associated with this trait, calving records from 2 time periods were used to estimate daughter averages for twinning for Holstein bulls. Multiple analyses were conducted and compared including GWAS, genomic prediction, and gene set enrichment analysis for pathway detection. Although pathway analysis did not yield many congruent pathways of interest between data sets, it did indicate two of interest. Both pathways have ties to the strong candidate region on BTA11 from the genome-wide association analysis across data sets. This region does not overlap with previously identified quantitative trait loci regions for twinning or ovulation rate in cattle. The strongest associated SNPs were upstream from 2 candidate genes LHCGR and FSHR, which are involved in folliculogenesis. Genomic prediction showed a moderate correlation accuracy (0.43) when predicting genomic breeding values for bulls with estimates from calving records from 2010 to 2016. Future analysis of the region on BTA11 and the relation of the candidate genes could improve this accuracy.     

Single-step genomic evaluation of milk production traits in Canadian Alpine and Saanen dairy goats

Genomic evaluations are routine in most plant and livestock breeding programs but are used infrequently in dairy goat breeding schemes. In this context, the purpose of this study was to investigate the use of the single-step genomic BLUP method for predicting genomic breeding values for milk production traits (milk, protein, and fat yields; protein and fat percentages) in Canadian Alpine and Saanen dairy goats. There were 6,409 and 12,236 Alpine records and 3,434 and 5,008 Saanen records for each trait in first and later lactations, respectively, and a total of 1,707 genotyped animals (833 Alpine and 874 Saanen). Two validation approaches were used, forward validation (i.e., animals born after 2013 with an average estimated breeding value accuracy from the full data set ≥0.50) and forward cross-validation (i.e., subsets of all animals included in the forward validation were used in successive replications). The forward cross-validation approach resulted in similar validation accuracies (0.55 to 0.66 versus 0.54 to 0.61) and biases (?0.01 to –0.07 versus ?0.03 to 0.11) to the forward validation when averaged across traits. Additionally, both single and multiple-breed analyses were compared, and similar average accuracies and biases were observed across traits. However, there was a small gain in accuracy from the use of multiple-breed models for the Saanen breed. A small gain in validation accuracy for genomically enhanced estimated breeding values (GEBV) relative to pedigree-based estimated breeding values (EBV) was observed across traits for the Alpine breed, but not for the Saanen breed, possibly due to limitations in the validation design, heritability of the traits evaluated, and size of the training populations. Trait-specific gains in theoretical accuracy of GEBV relative to EBV for the validation animals ranged from 17 to 31% in Alpine and 35 to 55% in Saanen, using the cross-validation approach. The GEBV predicted from the full data set were 12 to 16% more accurate than EBV for genotyped animals, but no gains were observed for nongenotyped animals. The largest gains were found for does without lactation records (35–41%) and bucks without daughter records (46–54%), and consequently, the implementation of genomic selection in the Canadian dairy goat population would be expected to increase selection accuracy for young breeding candidates. Overall, this study represents the first step toward implementation of genomic selection in Canadian dairy goat populations.     

Improving national fertility evaluations by accounting for the rapid rise of embryo transfer in US dairy cattle

Dairy producers have improved fertility of their herds by selecting bulls with higher conception rate evaluations. This research was motivated by the rapid increase in embryo transfer (ET) use to 11% of recent births and >1 million total births, with >5 times as many ET calves born in the United States in 2021 compared with just 5 yr earlier. Historical data used in genetic evaluations are stored in the National Cooperator Database. Recent records in the national pedigree database revealed that only 1% of ET calves have corresponding ET records in the breeding event database, 2% are incorrectly reported as artificial inseminations, and 97% have no associated breeding event. Embryo donation events are also rarely reported. Herd years reporting >10% of calves born by ET but less than half of the expected number of ET breeding events were removed to avoid potential biases. Heifer, cow, and sire conception rate evaluations were recalculated with this new data set according to the methods used for the official national evaluations. The edits removed about 1% of fertility records in the most recent 4 yr. Subsequent analysis showed that censoring herd years with inconsistent ET reporting had little effect on most bulls except for the highest ranking, younger bulls popular for ET use, and with largest effects on genomic selection. Improved ET reporting will be critical for providing accurate fertility evaluations, especially as the popularity of these advanced reproductive technologies continues to rise.     

Genetic and genomic analysis of age at first insemination in Israeli dairy cattle

We performed a genetic analysis of age at first insemination, including estimation of the heritability and genetic correlations with other economic traits, and the consequences of including this trait in the Israeli selection index. The genetic factors affecting age at first insemination were determined via GWAS. Five data sets were analyzed. Data sets 1, 2, and 3 were used to compute variance components among age at first insemination, first calving age, days from first insemination to calving, and the 9 traits included in the Israel breeding index. Heritabilities for age at first insemination, calving age, and days from first insemination to calving in Israeli Holsteins as computed by REML individual animal model analyses of 273,239 Israeli Holstein cows were 0.072, 0.042, and 0.014. The estimated genetic correlation between the first 2 traits was 0.88. In addition to the fact that heritability of age at first insemination is 1.7 times the heritability for calving, the former trait has the advantage that the number of records is greater, and the records are generated earlier. Absolute values of the genetic and residual correlations between age at first insemination and the 9 traits included in the Israeli index were all less than 0.2. Data set 4 included first insemination dates of 1,181,600 calves born from 1985 through 2018. Genetic evaluations were computed by a single trait animal model. Annual phenotypic and genetic trends for age at first calving for calves born since 1985 were “positive,” that is, economically negative, at 0.320 ± 0.003 and 0.169 ± 0.005 d, respectively. Applying the GCTA-GREML software, 54% of variance in the transmitting ability of 1,585 sires could be explained by considering all 40,498 markers included in the GWAS analysis. The significant markers were mainly associated with milk production genes. The SNP UA-IFASA-8854 on chromosome 11 had the lowest probability value, 1.2 × 10^?24. This marker is located between the genes RETSAT and ELMOD3, both of which are overexpressed in human mammary glands. The gene RETSAT is reported to be essential for lipid accumulation and adipogenesis promotion. Gene enrichment analysis found that genes in the genomic region flanking significant markers are associated with vasopressin receptor activity, which was shown to mediate puberty in humans. If age at first insemination is included in the index with a weighting to account for 9% of the index, reductions of 2.8 and 2.6 d for age at first insemination and first calving age after 10 yr of selection are predicted, as compared with reductions of 1.4 and 1.1 d with the current index. Gains for the other index traits are only marginally affected. We suggest selection on age at first insemination as an alternative to selection for early calving.