Investigating the benefits and perils of importing genetic material in small cattle breeding programs via simulation.

Small breeding programs are limited in achieving competitive genetic gain and prone to high rates of inbreeding. Thus, they often import genetic material to increase genetic gain and to limit the loss of genetic variability. However, the benefit of import depends on the strength of genotype-by-environment interaction. Import also diminishes the relevance of domestic selection and the use of domestic breeding animals. Introduction of genomic selection has potentially exacerbated this issue, but is also opening the potential for smaller breeding programs. The aim of this paper was to determine when and to what extent small breeding programs benefit from importing genetic material by quantifying the genetic gain as well as the sources of genetic gain. We simulated 2 cattle breeding programs of the same breed that represented a large foreign and a small domestic breeding program. The programs differed in selection parameters of sire selection, and in the initial genetic mean and annual genetic gain. We evaluated a control scenario without the use of foreign sires in the domestic breeding program and 24 scenarios that varied the percentage of domestic dams mated with foreign sires, the genetic correlation between the breeding programs (0.8 or 0.9), and the time of implementing genomic selection in the domestic compared with the foreign breeding program (concurrently or with a 10-yr delay). We compared the scenarios based on the genetic gain and genic standard deviation. Finally, we partitioned breeding values and genetic trends of the scenarios to quantify the contribution of domestic selection and import to the domestic genetic gain. The simulation revealed that when both breeding programs implemented genomic selection simultaneously, the use of foreign sires increased domestic genetic gain only when genetic correlation was 0.9 (10%-18% increase). In contrast, when the domestic breeding program implemented genomic selection with a 10-yr delay, import increased genetic gain at both tested correlations, 0.8 (5%-23% increase) and 0.9 (15%-53% increase). The increase was significant when we mated at least 10% or 25% domestic females with foreign sires and increased with the increasing use of foreign sires, but with a diminishing return. The partitioning analysis revealed that the contribution of import expectedly increased with the increased use of foreign sires. However, the increase did not depend on the genetic correlation and was not proportional to the increase in domestic genetic gain. This represents a peril for small breeding programs because they could be overly relying on import with diminishing returns for the genetic gain, marginal benefit for the genetic variability, and large loss of the domestic germplasm. The benefit and peril of import depends on an interplay of genetic correlation, extent of using foreign sires, and a breeding scheme. It is therefore crucial that small breeding programs assess the possible benefits of import beyond domestic selection. The benefit of import should be weighed against the perils of decreased use of domestic sires and decreased contribution and value of domestic selection.

Genomic selection for genotype performance and stability using information on multiple traits and multiple environments.

KEY MESSAGE: The inclusion of multiple traits and multiple environments within a partially separable factor analytic approach for genomic selection provides breeders with an informative framework to utilise genotype by environment by trait interaction for efficient selection. This paper develops a single-stage genomic selection (GS) approach which incorporates information on multiple traits and multiple environments within a partially separable factor analytic framework. The factor analytic linear mixed model is an effective method for analysing multi-environment trial (MET) datasets, but has not been extended to GS for multiple traits and multiple environments. The advantage of using all information is that breeders can utilise genotype by environment by trait interaction (GETI) to obtain more accurate predictions across correlated traits and environments. The partially separable factor analytic linear mixed model (SFA-LMM) developed in this paper is based on a three-way separable structure, which includes a factor analytic matrix between traits, a factor analytic matrix between environments and a genomic relationship matrix between genotypes. A diagonal matrix is then added to enable a different genotype by environment interaction (GEI) pattern for each trait and a different genotype by trait interaction (GTI) pattern for each environment. The results show that the SFA-LMM provides a better fit than separable approaches and a comparable fit to non-separable and partially separable approaches. The distinguishing feature of the SFA-LMM is that it will include fewer parameters than all other approaches as the number of genotypes, traits and environments increases. Lastly, a selection index is used to demonstrate simultaneous selection for overall performance and stability. This research represents an important continuation in the advancement of plant breeding analyses, particularly with the advent of high-throughput datasets involving a very large number of genotypes, traits and environments.

Association of inbreeding and regional equine leucocyte antigen homozygosity with the prevalence of insect bite hypersensitivity in Old Kladruber horse.

Inbreeding depression is the reduction of performance caused by mating of close relatives. In livestock populations, inbreeding depression has been traditionally estimated by regression of phenotypes on pedigree inbreeding coefficients. This estimation can be improved by utilising genomic inbreeding coefficients. Here we estimate inbreeding depression for insect bite hypersensitivity (IBH) prevalence, the most common allergic horse disease worldwide, in Old Kladruber horse. In a deep pedigree with 3214 horses (187 genotyped), we used a generalised linear mixed model with IBH phenotype from 558 horses examined between 1996 and 2009 (1368 records). In addition to the classical pedigree information, we used the single-step approach that enabled joint use of pedigree and genomic information to estimate inbreeding depression overall genome and equine leucocyte antigen (ELA) class II region. Significant inbreeding depression was observed in all models fitting overall inbreeding coefficients (odds ratio between 1.018 and 1.074, P < 0.05) with the exception of Kalinowski's new inbreeding (P = 0.0516). The increase of ELA class II inbreeding was significantly associated with increased prevalence of IBH (odds ratio 1.018; P = 0.027). However, when fitted jointly with the overall inbreeding coefficient, the effect of ELA class II inbreeding was not significant (odds ratio 1.016; P = 0.062). Overall, the higher ELA class II and/or overall inbreeding (pedigree or genomic) was associated with increased prevalence of IBH in Old Kladruber horses. The single-step approach provides an efficient use of all the available pedigree, genomic, and phenotype information for estimation of overall and regional inbreeding effects.

Genomic evaluation of Brown Swiss dairy cattle with limited national genotype data and integrated external information.

This study demonstrated the feasibility of a genomic evaluation for the dairy cattle population for which the small national training population can be complemented with foreign information from international evaluations. National test-day milk yield data records for the Slovenian Brown Swiss cattle population were analyzed. Genomic evaluation was carried out using the single-step genomic best linear unbiased prediction method (ssGBLUP), resulting in genomic estimated breeding values (GEBV). The predominantly female group of genotyped animals, representing the national training population in the single-step genomic evaluation, was further augmented with 7,024 genotypes of foreign progeny-tested sires from an international Brown Swiss InterGenomics genomic evaluation (https://interbull.org/ib/whole_cop). Additionally, the estimated breeding values for the altogether 7,246 genotyped domestic and foreign sires from the 2019 sire multiple across-country evaluation (MACE), were added to the ssGBLUP as external pseudophenotypic information. The ssGBLUP method, with integration of MACE information by avoiding double counting, was then performed, resulting in MACE-enhanced GEBV (GEBVM). The methods were empirically validated with forward prediction. The validation group consisted of 315 domestic males and 1,041 domestic females born after 2012. Increase, inflation, and bias of the GEBV(M) reliability (REL) were assessed for the validation group with a focus on females. All individuals in the validation benefited from genomic evaluations using both methods, but the GEBV(M) REL increased most for the youngest selection candidates. Up to 35 points of GEBV REL could be assigned to national genomic information, and up to 17 points of GEBVM REL could additionally be attributed to the integration of foreign sire genomic and MACE information. Results indicated that the combined foreign progeny-tested sire genomic and external MACE information can be used in the single-step genomic evaluation as an equivalent replacement for domestic phenotypic information. Thus, an equal or slightly higher genomic breeding value REL was obtained sooner than the pedigree-based breeding value REL for the female selection candidates. When the abundant foreign progeny-tested sire genomic and MACE information was used to complement available national genomic and phenotypic information in single-step genomic evaluation, the genomic breeding value REL for young-female selection candidates increased approximately 10 points. Use of international information provides the possibility to upgrade small national training populations and obtain satisfying reliability of genomic breeding values even for the youngest female selection candidates, which will help to increase selection efficiency in the future.

Efficient use of genomic information for sustainable genetic improvement in small cattle populations.

In this study, we compared genetic gain, genetic variation, and the efficiency of converting variation into gain under different genomic selection scenarios with truncation or optimum contribution selection in a small dairy population by simulation. Breeding programs have to maximize genetic gain but also ensure sustainability by maintaining genetic variation. Numerous studies have shown that genomic selection increases genetic gain. Although genomic selection is a well-established method, small populations still struggle with choosing the most sustainable strategy to adopt this type of selection. We developed a simulator of a dairy population and simulated a model after the Slovenian Brown Swiss population with ∼10,500 cows. We compared different truncation selection scenarios by varying (1) the method of sire selection and their use on cows or bull-dams, and (2) selection intensity and the number of years a sire is in use. Furthermore, we compared different optimum contribution selection scenarios with optimization of sire selection and their usage. We compared scenarios in terms of genetic gain, selection accuracy, generation interval, genetic and genic variance, rate of coancestry, effective population size, and conversion efficiency. The results showed that early use of genomically tested sires increased genetic gain compared with progeny testing, as expected from changes in selection accuracy and generation interval. A faster turnover of sires from year to year and higher intensity increased the genetic gain even further but increased the loss of genetic variation per year. Although maximizing intensity gave the lowest conversion efficiency, faster turnover of sires gave an intermediate conversion efficiency. The largest conversion efficiency was achieved with the simultaneous use of genomically and progeny-tested sires that were used over several years. Compared with truncation selection, optimizing sire selection and their usage increased the conversion efficiency by achieving either comparable genetic gain for a smaller loss of genetic variation or higher genetic gain for a comparable loss of genetic variation. Our results will help breeding organizations implement sustainable genomic selection.

National single-step genomic method that integrates multi-national genomic information.

The aim of this paper was to develop a national single-step genomic BLUP that integrates multi-national genomic estimated breeding values (EBV) and associated reliabilities without double counting dependent data contributions from the different evaluations. Simultaneous use of all data, including phenotypes, pedigree, and genotypes, is a condition to obtain unbiased EBV. However, this condition is not always fully met, mainly due to unavailability of foreign raw data for imported animals. In dairy cattle genetic evaluations, this issue is traditionally tackled through the multiple across-country evaluation (MACE) of sires, performed by Interbull Centre (Uppsala, Sweden). Multiple across-country evaluation regresses all the available national information onto a joint pedigree to obtain country-specific rankings of all sires without sharing the raw data. In the context of genomic selection, the issue is handled by exchanging sire genotypes and by using MACE information (i.e., MACE EBV and reliabilities), as a valuable source of "phenotypic" data. Although all the available data are considered, these "multi-national" genomic evaluations use multi-step methods assuming independence of various sources of information, which is not met in all situations. We developed a method that handles this by single-step genomic evaluation that jointly (1) uses national phenotypic, genomic, and pedigree data; (2) uses multi-national genomic information; and (3) avoids double counting dependent data contributions from an animal's own records and relatives' records. The method was demonstrated by integrating multi-national genomic EBV and reliabilities of Brown Swiss sires, included in the InterGenomics consortium at Interbull Centre, into the national evaluation in Slovenia. The results showed that the method could (1) increase reliability of a national (genomic) evaluation; (2) provide consistent ranking of all animals: bulls, cows, and young animals; and (3) increase the size of a genomic training population. These features provide more efficient and transparent selection throughout a breeding program.

Partition of genetic trends by origin in Landrace and Large-White pigs.

The objective of this study was to analyse the effectiveness of genetic improvement via domestic selection and import for backfat thickness and time on test in a conventional pig breeding programme for Landrace (L) and Large-White (LW) breeds. Phenotype data was available for 25 553 L and 10 432 LW pigs born between 2002 and 2012 from four large-scale farms and 72 family farms. Pedigree information indicated whether each animal was born and registered within the domestic breeding programme or has been imported. This information was used for defining the genetic groups of unknown parents in a pedigree and the partitioning analysis. Breeding values were estimated using a Bayesian analysis of an animal model with and without genetic groups. Such analysis enabled full Bayesian inference of the genetic trends and their partitioning by the origin of germplasm. Estimates of genetic group indicated that imported germplasm was overall better than domestic and substantial changes in estimates of breeding values was observed when genetic group were fitted. The estimated genetic trends in L were favourable and significantly different from zero by the end of the analysed period. Overall, the genetic trends in LW were not different from zero. The relative contribution of imported germplasm to genetic trends was large, especially towards the end of analysed period with 78% and 67% in L and from 50% to 67% in LW. The analyses suggest that domestic breeding activities and sources of imported animals need to be re-evaluated, in particular in LW breed.

Comparison between sire-maternal grandsire and animal models for genetic evaluation of longevity in a dairy cattle population with small herds.

Survival analysis techniques for sire-maternal grandsire (MGS) and animal models were used to test the genetic evaluation of longevity in a Slovenian Brown cattle population characterized by small herds. Three genetic models were compared: a sire-MGS model for bulls and an approximate animal model based on estimated breeding values (EBV) from the sire-MGS model for cows, an animal model, and an animal model based on the estimated variance components from the sire-MGS model. In addition, modeling the contemporary group effect was defined as either a herd or a herd-year (HY) effect. With various restrictions on the minimum HY group size (from 1 to 10 cows per HY), changes in estimates of variance components, and consequently also in EBV, were observed for the sire-MGS and animal models. Variance of contemporary group effects decreased when an HY effect was fitted instead of a herd effect. In the case of a sire-MGS model, estimates of additive genetic variance were mostly robust to changes in minimum HY group size or fitting herd or HY effect, whereas they increased in the animal model when HY instead of herd effects was fitted, possibly revealing some confounding between cow EBV and contemporary group effect. Estimated heritabilities from sire-MGS models were between 0.091 and 0.119 and were mainly influenced by the restriction on the HY group size. Estimated heritabilities from animal models were higher: between 0.125 and 0.160 when herd effect was fitted and between 0.171 and 0.210 when HY effect was fitted. Rank correlations between the animal model and the approximate animal model based on EBV from the sire-MGS model were high: 0.94 for cows and 0.93 for sires when a herd effect was fitted and 0.90 for cows and 0.93 for sires when an HY effect was fitted. Validation performed on the independent validation data set revealed that the correlation between sire EBV and daughter survival were slightly higher with the approximate animal model based on EBV from the sire-MGS model compared with the animal model. The correlations between the sire EBV and daughter survival were higher when the model included an HY effect instead of a herd effect. To avoid confounding and reduce computational requirements, it is suggested that the approximate animal model based on EBV from the sire-MGS model and HY as a contemporary group effect is an interesting compromise for practical applications of genetic evaluation of longevity in cattle populations.

Genomic evaluations using similarity between haplotypes.

Long-range phasing and haplotype library imputation methodologies are accurate and efficient methods to provide haplotype information that could be used in prediction of breeding value or phenotype. Modelling long haplotypes as independent effects in genomic prediction would be inefficient due to the many effects that need to be estimated and phasing errors, even if relatively low in frequency, exacerbate this problem. One approach to overcome this is to use similarity between haplotypes to model covariance of genomic effects by region or of animal breeding values. We developed a simple method to do this and tested impact on genomic prediction by simulation. Results show that the diagonal and off-diagonal elements of a genomic relationship matrix constructed using the haplotype similarity method had higher correlations with the true relationship between pairs of individuals than genomic relationship matrices built using unphased genotypes or assumed unrelated haplotypes. However, the prediction accuracy of such haplotype-based prediction methods was not higher than those based on unphased genotype information.

The contribution of dominance and inbreeding depression in estimating variance components for litter size in Pannon White rabbits.

In a synthetic closed population of Pannon White rabbits, additive (VA ), dominance (VD ) and permanent environmental (VPe ) variance components as well as doe (bF d ) and litter (bF l ) inbreeding depression were estimated for the number of kits born alive (NBA), number of kits born dead (NBD) and total number of kits born (TNB). The data set consisted of 18,398 kindling records of 3883 does collected from 1992 to 2009. Six models were used to estimate dominance and inbreeding effects. The most complete model estimated VA and VD to contribute 5.5 ± 1.1% and 4.8 ± 2.4%, respectively, to total phenotypic variance (VP ) for NBA; the corresponding values for NBD were 1.9 ± 0.6% and 5.3 ± 2.4%, for TNB, 6.2 ± 1.0% and 8.1 ± 3.2% respectively. These results indicate the presence of considerable VD . Including dominance in the model generally reduced VA and VPe estimates, and had only a very small effect on inbreeding depression estimates. Including inbreeding covariates did not affect estimates of any variance component. A 10% increase in doe inbreeding significantly increased NBD (bF d  = 0.18 ± 0.07), while a 10% increase in litter inbreeding significantly reduced NBA (bF l  = -0.41 ± 0.11) and TNB (bF l  = -0.34 ± 0.10). These findings argue for including dominance effects in models of litter size traits in populations that exhibit significant dominance relationships.

Effects of two probiotic additives containing Bacillus spores on carcass characteristics, blood lipids and cecal volatile fatty acids in meat type chickens.

The objective of this study was to evaluate effects of two commercially available probiotic additives, containing Bacillus spores, on carcass and meat characteristics, serum lipids and concentration of cecal volatile fatty acids of meat type chickens. Birds were fed regular corn-soy meal based feed (control), supplemented with additive A, containing 1.6 × 10(6) spores per gram of feed of Bacillus subtilis and Bacillus licheniformis (group A) or additive B, containing the same concentration of Bacillus cereus var. toyoi spores (group B). One hundred and twenty birds (20 per replicate) were slaughtered at the age of 55 days. Results showed that birds in group B had higher (p < 0.05) final body weight compared to birds from group A and higher carcass weights and yield percentages compared with control. Breasts and whole legs were also heavier in group B, compared to control, but not the yield. Group A had higher yield of wings and lower abdominal fat weight compared to group B (p< 0.05), but not compared with control. Total cholesterol was not affected by the dietary treatment, on contrary both probiotics elevated the LDL (p < 0.05) and lowered HDL cholesterol, thus unfavourably changed animal's blood serum cholesterol profile. Both probiotics influenced the cecal fermentation, which was observed as decrease in cecal concentrations of propionic, butyric, n-butyric and n-valeric acids, but the differences compared to control group were statistically significant for group A only. It was established that probiotic additive B was more effective regarding carcass and meat part weights than additive A, however the animals from group B also had more abdominal fat and their meat had significantly higher conductivity than control group, which is not considered as beneficial.

Porous concrete block as an environmental enrichment device increases activity of laying hens in cages.

The purpose of this study was to consider the influence of simple and cheap environmental enrichment such as porous concrete on the behavior of laying hens in conventional cages. Forty brown laying hens were housed in individual wire mesh cages: 20 in experimental cages with porous concrete block provided for pecking and 20 in a control group without concrete block provided. Porous concrete block (5 cm length x 5 cm width x 5 cm height) was mounted on the side wall at the height of the hen's head. Behavior was studied from 42 to 48 wk of age. A group of 8 hens was filmed for 24 h, and the camera was moved each day so that all 40 hens were recorded over 5 d each wk. Videotaping was performed in wk 1, 3, 5, and 7 of the experiment. States (long-term behavior) were observed with 5-min interval recording (feeding, preening, resting, and remaining inactive), whereas events (short-term activities) were observed with instantaneous recording (drinking, pecking concrete, pecking neighbors, pecking cage, and attempting to escape). Data were analyzed with generalized linear mixed model with binomial distribution for states, and Poisson distribution for events. Monte Carlo Markov Chain methods were used to estimate model parameters. Because posterior distributions of quantities of interest were skewed, medians and standard errors are reported. Hens in experimental cages were more active in long-term behavior than controls (64.9 +/- 1.9 and 59.3 +/- 1.9% of the light period, respectively). Correspondingly, hens in the control group showed more long-term inactivity. In addition to pecking the porous concrete block, hens in experimental cages also showed other short-term activities with greater frequency (4.10 +/- 0.31 and 3.51 +/- 0.25 events per h, respectively). Our hypothesis that hens in enriched cages would have a greater level of activity was confirmed. Provision of a piece of porous concrete block as a pecking substrate enriched the environment of the birds at negligible cost.

Genotoxicity evaluation of water soil leachates by Ames test, comet assay, and preliminary Tradescantia micronucleus assay.

Combining genotoxicity/mutagenicity tests and physico-chemical methodologies can be useful for determining the potential genotoxic contaminants in soil samples. The aim of our study was to evaluate the genotoxicity of soil by applying an integrated physico-chemical-biological approach. Soil samples were collected at six sampling points in a Slovenian industrial and agricultural region where contamination by heavy metals and sulphur dioxide (SO(2)) are primarily caused by a nearby power plant. The in vitro alkaline version of the comet assay on water soil leachates was performed with Caco-2 and HepG2 cells. A parallel genotoxicity evaluation of the samples was performed by Ames test using Salmonella typhimurium and the Tradescantia micronucleus test. Pedological analyses, heavy metal content determination, and different physico-chemical analyses, were also performed utilizing standard methodology. Water leachates of soil samples were prepared according to standard methods. Since only a battery of biotests with prokaryotic and eukaryotic organisms or cells can accurately estimate the effects of (geno)toxicants in soil samples and water soil leachates, a combination of three bioassays, with cells or organisms belonging to different trophic levels, was used. Genotoxicity of all six water soil leachates was proven by the comet assay on both human cell lines, however no positive results were detected by bacterial assay, Ames test. The Tradescantia micronucleus assay showed increase in micronuclei formation for three samples. According to these results we can assume that the comet assay was the most sensitive assay, followed by the micronucleus test. The Ames test does not appear to be sensitive enough for water soil leachates genotoxicity evaluations where heavy metal contamination is anticipated.