Simulation of functional additive and non-additive genetic effects using statistical estimates from quantitative genetic models.

Stochastic simulation software is commonly used to aid breeders designing cost-effective breeding programs and to validate statistical models used in genetic evaluation. An essential feature of the software is the ability to simulate populations with desired genetic and non-genetic parameters. However, this feature often fails when non-additive effects due to dominance or epistasis are modeled, as the desired properties of simulated populations are estimated from classical quantitative genetic statistical models formulated at the population level. The software simulates underlying functional effects for genotypic values at the individual level, which are not necessarily the same as effects from statistical models in which dominance and epistasis are included. This paper provides the theoretical basis and mathematical formulas for the transformation between functional and statistical effects in such simulations. The transformation is demonstrated with two statistical models analyzing individual phenotypes in a single population (common in animal breeding) and plot phenotypes of three-way hybrids involving two inbred populations (observed in some crop breeding programs). We also describe different methods for the simulation of functional effects for additive genetics, dominance, and epistasis to achieve the desired levels of variance components in classical statistical models used in quantitative genetics.

The soil microbiome of Lolium perenne L. depends on host genotype, is modified by nitrogen level and varies across season.

Genotype by environment interactions (G × E) are frequently observed in herbage production. Understanding the underlying biological mechanisms is important for achieving stable and predictive outputs across production environments. The microbiome is gaining increasing attention as a significant contributing factor to G × E. Here, we focused on the soil microbiome of perennial ryegrass (Lolium perenne L.) grown under field conditions and investigated the soil microbiome variation across different ryegrass varieties to assess whether environmental factors, such as seasonality and nitrogen levels, affect the microbial community. We identified bacteria, archaea, and fungi operational taxonomic units (OTUs) and showed that seasonality and ryegrass variety were the two factors explaining the largest fraction of the soil microbiome diversity. The strong and significant variety-by-treatment-by-seasonal cut interaction for ryegrass dry matter was associated with the number of unique OTUs within each sample. We identified seven OTUs associated with ryegrass dry matter variation. An OTU belonging to the Solirubrobacterales (Thermoleophilales) order was associated with increased plant biomass, supporting the possibility of developing engineered microbiomes for increased plant yield. Our results indicate the importance of incorporating different layers of biological data, such as genomic and soil microbiome data to improve the prediction accuracy of plant phenotypes grown across heterogeneous environments.

Scald resistance in hybrid rye (Secale cereale): genomic prediction and GWAS.

Rye (Secale cereale L.) is an important cereal crop used for food, beverages, and feed, especially in North-Eastern Europe. While rye is generally more tolerant to biotic and abiotic stresses than other cereals, it still can be infected by several diseases, including scald caused by Rhynchosporium secalis. The aims of this study were to investigate the genetic architecture of scald resistance, to identify genetic markers associated with scald resistance, which could be used in breeding of hybrid rye and to develop a model for genomic prediction for scald resistance. Four datasets with records of scald resistance on a population of 251 hybrid winter rye lines grown in 2 years and at 3 locations were used for this study. Four genomic models were used to obtain variance components and heritabilities of scald resistance. All genomic models included additive genetic effects of the parental components of the hybrids and three of the models included additive-by-additive epistasis and/or dominance effects. All models showed moderate to high broad sense heritabilities in the range of 0.31 (SE 0.05) to 0.76 (0.02). The model without non-additive genetic effects and the model with dominance effects had moderate narrow sense heritabilities ranging from 0.24 (0.06) to 0.55 (0.08). None of the models detected significant non-additive genomic variances, likely due to a limited data size. A genome wide association study was conducted to identify markers associated with scald resistance in hybrid winter rye. In three datasets, the study identified a total of twelve markers as being significantly associated with scald resistance. Only one marker was associated with a major quantitative trait locus (QTL) influencing scald resistance. This marker explained 11-12% of the phenotypic variance in two locations. Evidence of genotype-by-environment interactions was found for scald resistance between one location and the other two locations, which suggested that scald resistance was influenced by different QTLs in different environments. Based on the results of the genomic prediction models and GWAS, scald resistance seems to be a quantitative trait controlled by many minor QTL and one major QTL, and to be influenced by genotype-by-environment interactions.

Genomic prediction for root and yield traits of barley under a water availability gradient: a case study comparing different spatial adjustments.

BACKGROUND: In drought periods, water use efficiency depends on the capacity of roots to extract water from deep soil. A semi-field phenotyping facility (RadiMax) was used to investigate above-ground and root traits in spring barley when grown under a water availability gradient. Above-ground traits included grain yield, grain protein concentration, grain nitrogen removal, and thousand kernel weight. Root traits were obtained through digital images measuring the root length at different depths. Two nearest-neighbor adjustments (M1 and M2) to model spatial variation were used for genetic parameter estimation and genomic prediction (GP). M1 and M2 used (co)variance structures and differed in the distance function to calculate between-neighbor correlations. M2 was the most developed adjustment, as accounted by the Euclidean distance between neighbors. RESULTS: The estimated heritabilities ([Formula: see text]) ranged from low to medium for root and above-ground traits. The genetic coefficient of variation ([Formula: see text]) ranged from 3.2 to 7.0% for above-ground and 4.7 to 10.4% for root traits, indicating good breeding potential for the measured traits. The highest [Formula: see text] observed for root traits revealed that significant genetic change in root development can be achieved through selection. We studied the genotype-by-water availability interaction, but no relevant interaction effects were detected. GP was assessed using leave-one-line-out (LOO) cross-validation. The predictive ability (PA) estimated as the correlation between phenotypes corrected by fixed effects and genomic estimated breeding values ranged from 0.33 to 0.49 for above-ground and 0.15 to 0.27 for root traits, and no substantial variance inflation in predicted genetic effects was observed. Significant differences in PA were observed in favor of M2. CONCLUSIONS: The significant [Formula: see text] and the accurate prediction of breeding values for above-ground and root traits revealed that developing genetically superior barley lines with improved root systems is possible. In addition, we found significant spatial variation in the experiment, highlighting the relevance of correctly accounting for spatial effects in statistical models. In this sense, the proposed nearest-neighbor adjustments are flexible approaches in terms of assumptions that can be useful for semi-field or field experiments.

Metabolomic-genomic prediction can improve prediction accuracy of breeding values for malting quality traits in barley.

BACKGROUND: Metabolomics measures an intermediate stage between genotype and phenotype, and may therefore be useful for breeding. Our objectives were to investigate genetic parameters and accuracies of predicted breeding values for malting quality (MQ) traits when integrating both genomic and metabolomic information. In total, 2430 plots of 562 malting spring barley lines from three years and two locations were included. Five MQ traits were measured in wort produced from each plot. Metabolomic features used were 24,018 nuclear magnetic resonance intensities measured on each wort sample. Methods for statistical analyses were genomic best linear unbiased prediction (GBLUP) and metabolomic-genomic best linear unbiased prediction (MGBLUP). Accuracies of predicted breeding values were compared using two cross-validation strategies: leave-one-year-out (LOYO) and leave-one-line-out (LOLO), and the increase in accuracy from the successive inclusion of first, metabolomic data on the lines in the validation population (VP), and second, both metabolomic data and phenotypes on the lines in the VP, was investigated using the linear regression (LR) method. RESULTS: For all traits, we saw that the metabolome-mediated heritability was substantial. Cross-validation results showed that, in general, prediction accuracies from MGBLUP and GBLUP were similar when phenotypes and metabolomic data were recorded on the same plots. Results from the LR method showed that for all traits, except one, accuracy of MGBLUP increased when including metabolomic data on the lines of the VP, and further increased when including also phenotypes. However, in general the increase in accuracy of MGBLUP when including both metabolomic data and phenotypes on lines of the VP was similar to the increase in accuracy of GBLUP when including phenotypes on the lines of the VP. Therefore, we found that, when metabolomic data were included on the lines of the VP, accuracies substantially increased for lines without phenotypic records, but they did not increase much when phenotypes were already known. CONCLUSIONS: MGBLUP is a useful approach to combine phenotypic, genomic and metabolomic data for predicting breeding values for MQ traits. We believe that our results have significant implications for practical breeding of barley and potentially many other species.

Estimation and consequences of direct-maternal genetic and environmental covariances in models for genetic evaluation in broilers.

BACKGROUND: Maternal effects influence juvenile traits such as body weight and early growth in broilers. Ignoring significant maternal effects leads to reduced accuracy and inflated predicted breeding values. Including genetic and environmental direct-maternal covariances into prediction models in broilers can increase the accuracy and limit inflation of predicted breeding values better than simply adding maternal effects to the model. To test this hypothesis, we applied a model accounting for direct-maternal genetic covariance and direct-maternal environmental covariance to estimate breeding values. RESULTS: This model, and simplified versions of it, were tested using simulated broiler populations and then was applied to a large broiler population for validation. The real population analyzed consisted of a commercial line of broilers, for which body weight at a common slaughter age was recorded for 41 selection rounds. The direct-maternal genetic covariance was negative whereas the direct-maternal environmental covariance was positive. Simulated populations were created to mimic the real population. The predictive ability of the models was assessed by cross-validation, where the validation birds were all from the last five selection rounds. Accuracy of prediction was defined as the correlation between the predicted breeding values estimated without the phenotypic records of the validation population and a predictor. The predictors were the breeding values estimated using all the phenotypic information and the phenotypes corrected for the fixed effects, and for the simulated data, the true breeding values. In the real data, adding the environmental covariance, with or without also adding the genetic covariance, increased the accuracy, or reduced deflation of breeding values compared with a model not including dam-offspring covariance. Nevertheless, in the simulated data, reduction in the inflation of breeding values was possible and was associated with a gain in accuracy of up to 6% compared with a model not including both forms of direct-maternal covariance. CONCLUSIONS: In this paper, we propose a simple approach to estimate the environmental direct-maternal covariance using standard software for REML analysis. The genetic covariance between dam and offspring was negative whereas the corresponding environmental covariance was positive. Considering both covariances in models for genetic evaluation increased the accuracy of predicted breeding values.

Prediction of additive, epistatic, and dominance effects using models accounting for incomplete inbreeding in parental lines of hybrid rye and sugar beet.

Genomic models for prediction of additive and non-additive effects within and across different heterotic groups are lacking for breeding of hybrid crops. In this study, genomic prediction models accounting for incomplete inbreeding in parental lines from two different heterotic groups were developed and evaluated. The models can be used for prediction of general combining ability (GCA) of parental lines from each heterotic group as well as specific combining ability (SCA) of all realized and potential crosses. Here, GCA was estimated as the sum of additive genetic effects and within-group epistasis due to high degree of inbreeding in parental lines. SCA was estimated as the sum of across-group epistasis and dominance effects. Three models were compared. In model 1, it was assumed that each hybrid was produced from two completely inbred parental lines. Model 1 was extended to include three-way hybrids from parental lines with arbitrary levels of inbreeding: In model 2, parents of the three-way hybrids could have any levels of inbreeding, while the grandparents of the maternal parent were assumed completely inbred. In model 3, all parental components could have any levels of inbreeding. Data from commercial breeding programs for hybrid rye and sugar beet was used to evaluate the models. The traits grain yield and root yield were analyzed for rye and sugar beet, respectively. Additive genetic variances were larger than epistatic and dominance variances. The models' predictive abilities for total genetic value, for GCA of each parental line and for SCA were evaluated based on different cross-validation strategies. Predictive abilities were highest for total genetic values and lowest for SCA. Predictive abilities for SCA and for GCA of maternal lines were higher for model 2 and model 3 than for model 1. The implementation of the genomic prediction models in hybrid breeding programs can potentially lead to increased genetic gain in two different ways: I) by facilitating the selection of crossing parents with high GCA within heterotic groups and II) by prediction of SCA of all realized and potential combinations of parental lines to produce hybrids with high total genetic values.

Integrating a growth degree-days based reaction norm methodology and multi-trait modeling for genomic prediction in wheat.

Multi-trait and multi-environment analyses can improve genomic prediction by exploiting between-trait correlations and genotype-by-environment interactions. In the context of reaction norm models, genotype-by-environment interactions can be described as functions of high-dimensional sets of markers and environmental covariates. However, comprehensive multi-trait reaction norm models accounting for marker × environmental covariates interactions are lacking. In this article, we propose to extend a reaction norm model incorporating genotype-by-environment interactions through (co)variance structures of markers and environmental covariates to a multi-trait reaction norm case. To do that, we propose a novel methodology for characterizing the environment at different growth stages based on growth degree-days (GDD). The proposed models were evaluated by variance components estimation and predictive performance for winter wheat grain yield and protein content in a set of 2,015 F6-lines. Cross-validation analyses were performed using leave-one-year-location-out (CV1) and leave-one-breeding-cycle-out (CV2) strategies. The modeling of genomic [SNPs] × environmental covariates interactions significantly improved predictive ability and reduced the variance inflation of predicted genetic values for grain yield and protein content in both cross-validation schemes. Trait-assisted genomic prediction was carried out for multi-trait models, and it significantly enhanced predictive ability and reduced variance inflation in all scenarios. The genotype by environment interaction modeling via genomic [SNPs] × environmental covariates interactions, combined with trait-assisted genomic prediction, boosted the benefits in predictive performance. The proposed multi-trait reaction norm methodology is a comprehensive approach that allows capitalizing on the benefits of multi-trait models accounting for between-trait correlations and reaction norm models exploiting high-dimensional genomic and environmental information.

Relative importance of genotype, gene expression, and DNA methylation on complex traits in perennial ryegrass.

The growing demand for food and feed crops in the world because of growing population and more extreme weather events requires high-yielding and resilient crops. Many agriculturally important traits are polygenic, controlled by multiple regulatory layers, and with a strong interaction with the environment. In this study, 120 F2 families of perennial ryegrass (Lolium perenne L.) were grown across a water gradient in a semifield facility with subsoil irrigation. Genomic (single-nucleotide polymorphism [SNP]), transcriptomic (gene expression [GE]), and DNA methylomic (MET) data were integrated with feed quality trait data collected from control and drought sections in the semifield facility, providing a treatment effect. Deep root length (DRL) below 110 cm was assessed with convolutional neural network image analysis. Bayesian prediction models were used to partition phenotypic variance into its components and evaluated the proportion of phenotypic variance in all traits captured by different regulatory layers (SNP, GE, and MET). The spatial effects and effects of SNP, GE, MET, the interaction between GE and MET (GE × MET) and GE × treatment (GEControl and GEDrought ) interaction were investigated. Gene expression explained a substantial part of the genetic and spatial variance for all the investigated phenotypes, whereas MET explained residual variance not accounted for by SNPs or GE. For DRL, MET also contributed to explaining spatial variance. The study provides a statistically elegant analytical paradigm that integrates genomic, transcriptomic, and MET information to understand the regulatory mechanisms of polygenic effects for complex traits.

Effective Selection for Lower Mortality in Organic Pigs through Selection for Total Number Born and Number of Dead Piglets.

Selection for the number of living pigs on day 11 (L11) aims to reduce piglet mortality and increase litter size simultaneously. This approach could be sub-optimal, especially for organic pig breeding. This study evaluated the effect of selecting for a trait by separating it into two traits. Genetic parameters for L11, the total number born (TNB), and the number of dead piglets at day 11 (D11) were estimated using data obtained from an organic pig population in Denmark. Based on these estimates, two alternative breeding schemes were simulated. Specifically, selection was made using: (1) a breeding goal with L11 only versus (2) a breeding goal with TNB and D11. Different weightings for TNB and D11 were tested. The simulations showed that selection using the first breeding scheme (L11) produced lower annual genetic gain (0.201) compared to the second (TNB and D11; 0.207). A sensitivity analysis showed that the second scheme performed better because it exploited differences in heritability, and accounted for genetic correlations between the two traits. When the second breeding scheme placed more emphasis on D11, D11 declined, whereas genetic gain for L11 remained high (0.190). In conclusion, selection for L11 could be optimized by separating it into two correlated traits with different heritability, reducing piglet mortality and enhancing L11.

Correction: Salimiyekta et al. Random Regression Model for Genetic Evaluation and Early Selection in the Iranian Holstein Population. Animals 2021, 11, 3492.

There was an error in the original publication [...].

Association Studies and Genomic Prediction for Genetic Improvements in Agriculture.

To feed the fast growing global population with sufficient food using limited global resources, it is urgent to develop and utilize cutting-edge technologies and improve efficiency of agricultural production. In this review, we specifically introduce the concepts, theories, methods, applications and future implications of association studies and predicting unknown genetic value or future phenotypic events using genomics in the area of breeding in agriculture. Genome wide association studies can identify the quantitative genetic loci associated with phenotypes of importance in agriculture, while genomic prediction utilizes individual genetic value to rank selection candidates to improve the next generation of plants or animals. These technologies and methods have improved the efficiency of genetic improvement programs for agricultural production via elite animal breeds and plant varieties. With the development of new data acquisition technologies, there will be more and more data collected from high-through-put technologies to assist agricultural breeding. It will be crucial to extract useful information among these large amounts of data and to face this challenge, more efficient algorithms need to be developed and utilized for analyzing these data. Such development will require knowledge from multiple disciplines of research.

Identification of Candidate Genes Regulating Carcass Depth and Hind Leg Circumference in Simmental Beef Cattle Using Illumina Bovine Beadchip and Next-Generation Sequencing Analyses.

Genome-wide association studies are a robust means of identifying candidate genes that regulate economically important traits in farm animals. The aim of this study is to identify single-nucleotide polymorphisms (SNPs) and candidate genes potentially related to carcass depth and hind leg circumference in Simmental beef cattle. We performed Illumina Bovine HD Beadchip (~670 k SNPs) and next-generation sequencing (~12 million imputed SNPs) analyses of data from 1252 beef cattle, to which we applied a linear mixed model. Using a statistical threshold (p = 0.05/number of SNPs identified) and adopting a false discovery rate (FDR), we identified many putative SNPs on different bovine chromosomes. We identified 12 candidate genes potentially annotated with the markers identified, including CDKAL1 and E2F3, related to myogenesis and skeletal muscle development. The identification of such genes in Simmental beef cattle will help breeders to understand and improve related traits, such as meat yield.

Metabolomic spectra for phenotypic prediction of malting quality in spring barley.

We investigated prediction of malting quality (MQ) phenotypes in different locations using metabolomic spectra, and compared the prediction ability of different models, and training population (TP) sizes. Data of five MQ traits was measured on 2667 individual plots of 564 malting spring barley lines from three years and two locations. A total of 24,018 metabolomic features (MFs) were measured on each wort sample. Two statistical models were used, a metabolomic best linear unbiased prediction (MBLUP) and a partial least squares regression (PLSR). Predictive ability within location and across locations were compared using cross-validation methods. For all traits, more than 90% of the total variance in MQ traits could be explained by MFs. The prediction accuracy increased with increasing TP size and stabilized when the TP size reached 1000. The optimal number of components considered in the PLSR models was 20. The accuracy using leave-one-line-out cross-validation ranged from 0.722 to 0.865 and using leave-one-location-out cross-validation from 0.517 to 0.817. In conclusion, the prediction accuracy of metabolomic prediction of MQ traits using MFs was high and MBLUP is better than PLSR if the training population is larger than 100. The results have significant implications for practical barley breeding for malting quality.

Genetic variation in piglet mortality in outdoor organic production systems.

Piglet mortality from farrowing to weaning is a major concern, especially in outdoor organic production systems. This issue might impair animal welfare and generate economic losses for the farmer. In particular, it is difficult to apply management tools that are commonly used for indoor pig production systems to organic or outdoor production systems. Genetics and breeding approaches might be used to improve piglet survival. However, knowledge remains limited on the genetic background underlying survival traits in organic pigs that are born and reared outdoors. Here, we investigated the mortality of piglets from farrowing to weaning in an outdoor organic pig population and suggested genetic strategies to reduce piglet mortality in this production system. The experiment included mortality records of piglets from farrowing to weaning (around 69 days of age). Pedigree-based threshold models were used to analyse the mortality traits of piglets at 0-3 days of age, 4-11 days, and 12 days to weaning. Stillborn piglets were included in the group of piglets that died at 0-3 days of age. We found that the mortality rate from farrowing to weaning was, on average, 19.2%. However, most piglet deaths (79.1%) occurred at 0-11 days of age. As the age of piglets increased, the direct heritability of piglet mortality rose from 0 to 0.04, whereas maternal heritability decreased from 0.03 to a non-significant value. Piglets with higher BW had a lower mortality rate. However, the genetic correlations between maternal effects on piglet mortality and piglet BW were not significant; thus, selection for piglets with higher BW at around 10 days of age, through improving maternal genetics, would not reduce piglet mortality. Piglet mortality increased from sows with increasing number of parities. Crossbreeding also reduced piglet mortality. In conclusion, selection focusing on sow genotype, the use of younger sows, and crossbreeding could contribute to maintain piglet mortality at lower levels in outdoor organic pig production systems.

The Effect of Using Organic or Conventional Sires on Genetic Gain in Organic Pigs: A Simulation Study.

Current organic pig-breeding programs use pigs from conventional breeding populations. However, there are considerable differences between conventional and organic production systems. This simulation study aims to evaluate how the organic pig sector could benefit from having an independent breeding program. Two organic pig-breeding programs were simulated: one used sires from a conventional breeding population (conventional sires), and the other used sires from an organic breeding population (organic sires). For maintaining the breeding population, the conventional population used a conventional breeding goal, whereas the organic population used an organic breeding goal. Four breeding goals were simulated: one conventional breeding goal, and three organic breeding goals. When conventional sires were used, genetic gain in the organic population followed the conventional breeding goal, even when an organic breeding goal was used to select conventional sires. When organic sires were used, genetic gain followed the organic breeding goal. From an economic point of view, using conventional sires for breeding organic pigs is best, but only if there are no genotype-by-environment interactions. However, these results show that from a biological standpoint, using conventional sires biologically adapts organic pigs for a conventional production system.

Improvement of genomic prediction in advanced wheat breeding lines by including additive-by-additive epistasis.

KEY MESSAGE: Including additive and additive-by-additive epistasis in a NOIA parametrization did not yield orthogonal partitioning of genetic variances, nevertheless, it improved predictive ability in a leave-one-out cross-validation for wheat grain yield. Additive-by-additive epistasis is the principal non-additive genetic effect in inbred wheat lines and is potentially useful for developing cultivars based on total genetic merit; nevertheless, its practical benefits have been highly debated. In this article, we aimed to (i) evaluate the performance of models including additive and additive-by-additive epistatic effects for variance components (VC) estimation of grain yield in a wheat-breeding population, and (ii) to investigate whether including additive-by-additive epistasis in genomic prediction enhance wheat grain yield predictive ability (PA). In total, 2060 sixth-generation (F6) lines from Nordic Seed A/S breeding company were phenotyped in 21 year-location combinations in Denmark, and genotyped using a 15 K-Illumina-BeadChip. Three models were used to estimate VC and heritability at plot level: (i) "I-model" (baseline), (ii) "I + GA-model", extending I-model with an additive genomic effect, and (iii) "I + GA + GAA-model", extending I + GA-model with an additive-by-additive genomic effects. The I + GA-model and I + GA + GAA-model were based on the Natural and Orthogonal Interactions Approach (NOIA) parametrization. The I + GA + GAA-model failed to achieve orthogonal partition of genetic variances, as revealed by a change in estimated additive variance of I + GA-model when epistasis was included in the I + GA + GAA-model. The PA was studied using leave-one-line-out and leave-one-breeding-cycle-out cross-validations. The I + GA + GAA-model increased PA significantly (16.5%) compared to the I + GA-model in leave-one-line-out cross-validation. However, the improvement due to including epistasis was not observed in leave-one-breeding-cycle-out cross-validation. We conclude that epistatic models can be useful to enhance predictions of total genetic merit. However, even though we used the NOIA parameterization, the variance partition into orthogonal genetic effects was not possible.

Genomic prediction for grain yield and micro-environmental sensitivity in winter wheat.

Individuals within a common environment experience variations due to unique and non-identifiable micro-environmental factors. Genetic sensitivity to micro-environmental variation (i.e. micro-environmental sensitivity) can be identified in residuals, and genotypes with lower micro-environmental sensitivity can show greater resilience towards environmental perturbations. Micro-environmental sensitivity has been studied in animals; however, research on this topic is limited in plants and lacking in wheat. In this article, we aimed to (i) quantify the influence of genetic variation on residual dispersion and the genetic correlation between genetic effects on (expressed) phenotypes and residual dispersion for wheat grain yield using a double hierarchical generalized linear model (DHGLM); and (ii) evaluate the predictive performance of the proposed DHGLM for prediction of additive genetic effects on (expressed) phenotypes and its residual dispersion. Analyses were based on 2,456 advanced breeding lines tested in replicated trials within and across different environments in Denmark and genotyped with a 15K SNP-Illumina-BeadChip. We found that micro-environmental sensitivity for grain yield is heritable, and there is potential for its reduction. The genetic correlation between additive effects on (expressed) phenotypes and dispersion was investigated, and we observed an intermediate correlation. From these results, we concluded that breeding for reduced micro-environmental sensitivity is possible and can be included within breeding objectives without compromising selection for increased yield. The predictive ability and variance inflation for predictions of the DHGLM and a linear mixed model allowing heteroscedasticity of residual variance in different environments (LMM-HET) were evaluated using leave-one-line-out cross-validation. The LMM-HET and DHGLM showed good and similar performance for predicting additive effects on (expressed) phenotypes. In addition, the accuracy of predicting genetic effects on residual dispersion was sufficient to allow genetic selection for resilience. Such findings suggests that DHGLM may be a good choice to increase grain yield and reduce its micro-environmental sensitivity.

Random Regression Model for Genetic Evaluation and Early Selection in the Iranian Holstein Population.

The objective of this study was to use a model to predict breeding values for sires and cows at an early stage of the first lactation of cows and progeny groups in the Iranian Holstein population to enable the early selection of sires. An additional objective was to estimate genetic and phenotypic parameters associated with this model. The accuracy of predicted breeding values was investigated using cross-validation based on sequential genetic evaluations emulating yearly evaluation runs. The data consisted of 2,166,925 test-day records from 456,712 cows calving between 1990 and 2015. (Co)-variance components and breeding values were estimated using a random regression test-day model and the average information (AI) restricted maximum likelihood method (REML). Legendre polynomial functions of order three were chosen to fit the additive genetic and permanent environmental effects, and a homogeneous residual variance was assumed throughout lactation. The lowest heritability of daily milk yield was estimated to be just under 0.14 in early lactation, and the highest heritability of daily milk yield was estimated to be 0.18 in mid-lactation. Cross-validation showed a highly positive correlation of predicted breeding values between consecutive yearly evaluations for both cows and sires. Correlation between predicted breeding values based only on records of early lactation (5-90 days) and records including late lactation (181-305 days) were 0.77-0.87 for cows and 0.81-0.94 for sires. These results show that we can select sires according to their daughters' early lactation information before they finish the first lactation. This can be used to decrease generation interval and to increase genetic gain in the Iranian Holstein population.