Breeding schemes with optimum-contribution selection or truncation selection for beef cattle destined for use on dairy females.

We tested the hypothesis that the size of a beef cattle population destined for use on dairy females is smaller under optimum-contribution selection (OCS) than under truncation selection (TRS) at the same genetic gain (ΔG) and the same rate of inbreeding (ΔF). We used stochastic simulation to estimate true ΔG realized at a 0.005 ΔF in breeding schemes with OCS or TRS. The schemes for the beef cattle population also differed in the number of purebred offspring per dam and the total number of purebred offspring per generation. Dams of the next generation were exclusively selected among the one-year-old heifers. All dams were donors for embryo transfer and produced a maximum of 5 or 10 offspring. The total number of purebred offspring per generation was: 400, 800, 1,600 or 4,000 calves, and it was used as a measure of population size. Rate of inbreeding was predicted and controlled using pedigree relationships. Each OCS (TRS) scheme was simulated for 10 discrete generations and replicated 100 (200) times. The OCS scheme and the TRS scheme with a maximum of 10 offspring per dam required approximately 783 and 1,257 purebred offspring per generation to realize a true ΔG of €14 and a ΔF of 0.005 per generation. Schemes with a maximum of 5 offspring per dam required more purebred offspring per generation to realize a similar true ΔG and a similar ΔF. Our results show that OCS and multiple ovulation and embryo transfer act on selection intensity through different mechanisms to achieve fewer selection candidates and fewer selected sires and dams than under TRS at the same ΔG and a fixed ΔF. Therefore, we advocate the use of a breeding scheme with OCS and multiple ovulation and embryo transfer for beef cattle destined for use on dairy females because it is favorable both from an economic perspective and a carbon footprint perspective.

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.

Short- and long-term consequences of collaboration between Northern European Red dairy and dual-purpose cattle.

In Northern European countries, a great variety of Red cattle populations exists which can be broadly categorized in two groups: specialized dairy and dual-purpose breeds. Collaboration between these breeds (i.e. the exchange of sires across breeds) can be beneficial but is limited so far. The aim of this study was to demonstrate and evaluate consequences of collaboration between Red breeds using stochastic simulations. Two breeding lines (dairy type and dual purpose) were simulated. As a special aspect of this study, differences in genetic levels of breeding traits (milk production, beef production, mastitis resistance, fertility, feed efficiency) have been taken into account. Various scenarios were investigated where across-breed selection was either restricted or allowed and with different correlations between breeding goals in the two lines. The results of this study were influenced by the different genetic levels in breeding traits only in the first years of simulation. In the long run, the breed differences did not affect the degree of collaboration between lines. When the correlation between breeding goals was close to unity, the selection of external bulls was highly beneficial in terms of genetic gain and total monetary gain. Additionally, the lowest rate of inbreeding was found in that case. With decreasing correlations between environments, degree of cooperation between lines rapidly terminated and lines operated individually. In last years of simulation, cooperation was only found when the correlation between breeding goals was close to unity. From a long-term perspective, the exchange of breeding sires across lines also caused negative effects. In the dual-purpose line, deterioration of genetic gain in mastitis resistance and fertility was observed. Additionally, breeding lines genetically converged, which decreased genetic diversity. Collectively, short-term benefits and long-term negative effects have to be reconciled if collaboration between Red breeds in Northern Europe is to be pursued.

Pre-selection against a lethal recessive allele in breeding schemes with optimum-contribution selection or truncation selection.

BACKGROUND: We tested the hypothesis that breeding schemes with a pre-selection step, in which carriers of a lethal recessive allele (LRA) were culled, and with optimum-contribution selection (OCS) reduce the frequency of a LRA, control rate of inbreeding, and realise as much genetic gain as breeding schemes without a pre-selection step. METHODS: We used stochastic simulation to estimate true genetic gain realised at a 0.01 rate of true inbreeding (ΔFtrue) by breeding schemes that combined one of four pre-selection strategies with one of three selection strategies. The four pre-selection strategies were: (1) no carriers culled, (2) male carriers culled, (3) female carriers culled, and (4) all carriers culled. Carrier-status was known prior to selection. The three selection strategies were: (1) OCS in which [Formula: see text] was predicted and controlled using pedigree relationships (POCS), (2) OCS in which [Formula: see text] was predicted and controlled using genomic relationships (GOCS), and (3) truncation selection of parents. All combinations of pre-selection strategies and selection strategies were tested for three starting frequencies of the LRA (0.05, 0.10, and 0.15) and two linkage statuses with the locus that has the LRA being on a chromosome with or without loci affecting the breeding goal trait. The breeding schemes were simulated for 10 discrete generations (t = 1, …, 10). In all breeding schemes, ΔFtrue was calibrated to be 0.01 per generation in generations t = 4, …, 10. Each breeding scheme was replicated 100 times. RESULTS: We found no significant difference in true genetic gain from generations t = 4, …, 10 between breeding schemes with or without pre-selection within selection strategy. POCS and GOCS schemes realised similar true genetic gains from generations t = 4, …, 10. POCS and GOCS schemes realised 12% more true genetic gain from generations t = 4, …, 10 than truncation selection schemes. CONCLUSIONS: We advocate for OCS schemes with pre-selection against the LRA that cause animal suffering and high costs. At LRA frequencies of 0.10 or lower, OCS schemes in which male carriers are culled reduce the frequency of LRA, control rate of inbreeding, and realise no significant reduction in true genetic gain compared to OCS schemes without pre-selection against LRA.

Strategies Using Genomic Selection to Increase Genetic Gain in Breeding Programs for Wheat.

Conventional wheat-breeding programs involve crossing parental lines and subsequent selfing of the offspring for several generations to obtain inbred lines. Such a breeding program takes more than 8 years to develop a variety. Although wheat-breeding programs have been running for many years, genetic gain has been limited. However, the use of genomic information as selection criterion can increase selection accuracy and that would contribute to increased genetic gain. The main objective of this study was to quantify the increase in genetic gain by implementing genomic selection in traditional wheat-breeding programs. In addition, we investigated the effect of genetic correlation between different traits on genetic gain. A stochastic simulation was used to evaluate wheat-breeding programs that run simultaneously for 25 years with phenotypic or genomic selection. Genetic gain and genetic variance of wheat-breeding program based on phenotypes was compared to the one with genomic selection. Genetic gain from the wheat-breeding program based on genomic estimated breeding values (GEBVs) has tripled compared to phenotypic selection. Genomic selection is a promising strategy for improving genetic gain in wheat-breeding programs.

Phenotypically Selective Genotyping Realizes More Genetic Gains in a Rainbow Trout Breeding Program in the Presence of Genotype-by-Environment Interactions.

Selective genotyping of phenotypically superior animals may lead to bias and less accurate genomic breeding values (GEBV). Performing selective genotyping based on phenotypes measured in the breeding environment (B) is not necessarily a good strategy when the aim of a breeding program is to improve animals' performance in the commercial environment (C). Our simulation study compared different genotyping strategies for selection candidates and for fish in C in a breeding program for rainbow trout in the presence of genotype-by-environment interactions when the program had limited genotyping resources and unregistered pedigrees of individuals. For the reference population, selective genotyping of top and bottom individuals in C based on phenotypes measured in C led to the highest genetic gains, followed by random genotyping and then selective genotyping of top individuals in C. For selection candidates, selective genotyping of top individuals in B based on phenotypes measured in B led to the highest genetic gains, followed by selective genotyping of top and bottom individuals and then random genotyping. Selective genotyping led to bias in predicting GEBV. However, in scenarios that used selective genotyping of top fish in B and random genotyping of fish in C, predictions of GEBV were unbiased, with genetic correlations of 0.2 and 0.5 between traits measured in B and C. Estimates of variance components were sensitive to genotyping strategy, with an overestimation of the variance with selective genotyping of top and bottom fish and an underestimation of the variance with selective genotyping of top fish. Unbiased estimates of variance components were obtained when fish in B and C were genotyped at random. In conclusion, we recommend phenotypic genotyping of top and bottom fish in C and top fish in B for the purpose of selecting breeding animals and random genotyping of individuals in B and C for the purpose of estimating variance components when a genomic breeding program for rainbow trout aims to improve animals' performance in C.

Genomic Breeding Programs Realize Larger Benefits by Cooperation in the Presence of Genotype × Environment Interaction Than Conventional Breeding Programs.

Genotype × environment interaction (G × E) is of increasing importance for dairy cattle breeders due to international multiple-environment selection of animals as well as the differentiation of production environments within countries. This theoretical simulation study tested the hypothesis that genomic selection (GS) breeding programs realize larger genetic benefits by cooperation in the presence of G × E than conventional pedigree-based selection (PS) breeding programs. We simulated two breeding programs each with their own cattle population and environment. Two populations had either equal or unequal population sizes. Selection of sires was done either across environments (cooperative) or within their own environment (independent). Four scenarios, (GS/PS) × (cooperative/independent), were performed. The genetic correlation (r g ) between the single breeding goal trait expressed in two environments was varied between 0.5 and 0.9. We compared scenarios for genetic gain, rate of inbreeding, proportion of selected external sires, and the split-point r g that is the lowest value of r g for long-term cooperation. Between two equal-sized populations, cooperative GS breeding programs achieved a maximum increase of 19.3% in genetic gain and a maximum reduction of 24.4% in rate of inbreeding compared to independent GS breeding programs. The increase in genetic gain and the reduction in rate of inbreeding realized by GS breeding programs with cooperation were respectively at maximum 9.7% and 24.7% higher than those realized by PS breeding programs with cooperation. Secondly, cooperative GS breeding programs allowed a slightly lower split-point r g than cooperative PS breeding programs (0.85∼0.875 vs ≥ 0.9). Between two unequal-sized populations, cooperative GS breeding programs realized higher increase in genetic gain and showed greater probability for long-term cooperation than cooperative PS breeding programs. Secondly, cooperation using GS were more beneficial to the small population while also beneficial but much less to the large population. In summary, by cooperation in the presence of G × E, GS breeding programs realize larger improvements in terms of the genetic gain and rate of inbreeding, and have greater possibility of long-term cooperation than conventional PS breeding programs. Therefore, we recommend cooperative GS breeding programs in situations with mild to moderate G × E, depending on the sizes of two populations.

Bias in estimates of variance components in populations undergoing genomic selection: a simulation study.

BACKGROUND: After the extensive implementation of genomic selection (GS), the choice of the statistical model and data used to estimate variance components (VCs) remains unclear. A primary concern is that VCs estimated from a traditional pedigree-based animal model (P-AM) will be biased due to ignoring the impact of GS. The objectives of this study were to examine the effects of GS on estimates of VC in the analysis of different sets of phenotypes and to investigate VC estimation using different methods. Data were simulated to resemble the Danish Jersey population. The simulation included three phases: (1) a historical phase; (2) 20 years of conventional breeding; and (3) 15 years of GS. The three scenarios based on different sets of phenotypes for VC estimation were as follows: (1) Pheno1: phenotypes from only the conventional phase (1-20 years); (2) Pheno1 + 2: phenotypes from both the conventional phase and GS phase (1-35 years); (3) Pheno2: phenotypes from only the GS phase (21-35 years). Single-step genomic BLUP (ssGBLUP), a single-step Bayesian regression model (ssBR), and P-AM were applied. Two base populations were defined: the first was the founder population referred to by the pedigree-based relationship (P-base); the second was the base population referred to by the current genotyped population (G-base). RESULTS: In general, both the ssGBLUP and ssBR models with all the phenotypic and genotypic information (Pheno1 + 2) yielded biased estimates of additive genetic variance compared to the P-base model. When the phenotypes from the conventional breeding phase were excluded (Pheno2), P-AM led to underestimation of the genetic variance of P-base. Compared to the VCs of G-base, when phenotypes from the conventional breeding phase (Pheno2) were ignored, the ssBR model yielded unbiased estimates of the total genetic variance and marker-based genetic variance, whereas the residual variance was overestimated. CONCLUSIONS: The results show that neither of the single-step models (ssGBLUP and ssBR) can precisely estimate the VCs for populations undergoing GS. Overall, the best solution for obtaining unbiased estimates of VCs is to use P-AM with phenotypes from the conventional phase or phenotypes from both the conventional and GS phases.

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

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

ADAM-Plant: A Software for Stochastic Simulations of Plant Breeding From Molecular to Phenotypic Level and From Simple Selection to Complex Speed Breeding Programs.

Making decisions on plant breeding programs require plant breeders to be able to test different breeding strategies by taking into account all the crucial factors affecting crop genetic improvement. Due to the complexity of the decisions, computer simulation serves as an important tool for researchers and plant breeders. This paper describes ADAM-plant, which is a computer software that models breeding schemes for self-pollinated and cross-pollinated crop plants using stochastic simulation. The program simulates a population of plants and traces the genetic changes in the population under different breeding scenarios. It takes into account different population structures, genomic models, selection (strategies and units) and crossing strategies. It also covers important features e.g., allowing users to perform genomic selection (GS) and speed breeding, simulate genotype-by-environment interactions using multiple trait approach, simulate parallel breeding cycles and consider plot sizes. In addition, the software can be used to simulate datasets produced from very complex breeding program in order to test new statistical methodology to analyze such data. As an example, three wheat-breeding strategies were simulated in the current study: (1) phenotypic selection, (2) GS, and (3) speed breeding with genomic information. The results indicate that the genetic gain can be doubled by GS compared to phenotypic selection and genetic gain can be further increased considerably by speed breeding. In conclusion, ADAM-plant is an important tool for comparing strategies for plant breeding and for estimating the effects of allocation of different resources to the breeding program. In the current study, it was used to compare different methodologies for utilizing genomic information in cereal breeding programs for selection of best-fit breeding strategy as per available resources.

Response to Selection in Finite Locus Models with Nonadditive Effects.

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

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

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

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

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

The advantage of factorial mating under selection is uncovered by deterministically predicted rates of inbreeding.

Rates of inbreeding (DeltaF) in selected populations were predicted using the framework of long-term genetic contributions and validated against stochastic simulations. Deterministic predictions decomposed DeltaF into four components due to: finite population size, directional selection, covariance of genetic contribution of mates, and deviation of variance of family size from that expected from a Poisson distribution. Factorial (FM) and hierarchical (HM) mating systems were compared under mass and sib-index selection. Prediction errors were in most cases for DeltaF less than 10% and for rate of gain less than 5%. DelatF was higher with index than mass selection. DeltaF was lower with FM than HM in all cases except random selection. FM reduced the variance of the average breeding value of the mates of an individual. This reduced the impact of the covariance of contributions of mates on DeltaF. Thus, contributions of mates were less correlated with FM than HM, causing smaller deviations of converged contributions from the optimum contributions. With index selection, FM also caused a smaller variance of number of offspring selected from each parent. This reduced variance of family size reduced DeltaF further. FM increases the flexibility in breeding schemes for achieving the optimum genetic contributions.

Precision of methods for calculating identity-by-descent matrices using multiple markers.

A rapid, deterministic method (DET) based on a recursive algorithm and a stochastic method based on Markov Chain Monte Carlo (MCMC) for calculating identity-by-descent (IBD) matrices conditional on multiple markers were compared using stochastic simulation. Precision was measured by the mean squared error (MSE) of the relationship coefficients in predicting the true IBD relationships, relative to MSE obtained from using pedigree only. Comparisons were made when varying marker density, allele numbers, allele frequencies, and the size of full-sib families. The precision of DET was 75-99% relative to MCMC, but was not simply related to the informativeness of individual loci. For situations mimicking microsatellite markers or dense SNP, the precision of DET was > or = 95% relative to MCMC. Relative precision declined for the SNP, but not microsatellites as marker density decreased. Full-sib family size did not affect the precision. The methods were tested in interval mapping and marker assisted selection, and the performance was very largely determined by the MSE. A multi-locus information index considering the type, number, and position of markers was developed to assess precision. It showed a marked empirical relationship with the observed precision for DET and MCMC and explained the complex relationship between relative precision and the informativeness of individual loci.