Nonparallel genome changes within subpopulations over time contributed to genetic diversity within the US Holstein population.

Maintaining genetic variation in a population is important for long-term genetic gain. The existence of subpopulations within a breed helps maintain genetic variation and diversity. The 20,990 genotyped animals, representing the breeding animals in the year 2014, were identified as the sires of animals born after 2010 with at least 25 progenies, and females measured for type traits within the last 2 yr of data. K-means clustering with 5 clusters (C1, C2, C3, C4, and C5) was applied to the genomic relationship matrix based on 58,990 SNP markers to stratify the selected candidates into subpopulations. The general higher inbreeding resulting from within-cluster mating than across-cluster mating suggests the successful stratification into genetically different groups. The largest cluster (C4) contained animals that were less related to each animal within and across clusters. The average fixation index was 0.03, indicating that the populations were differentiated, and allele differences across the subpopulations were not due to drift alone. Starting with the selected candidates within each cluster, a family unit was identified by tracing back through the pedigree, identifying the genotyped ancestors, and assigning them to a pseudogeneration. Each of the 5 families (F1, F2, F3, F4, and F5) was traced back for 10 generations, allowing for changes in frequency of individual SNPs over time to be observed, which we call allele frequencies change. Alternative procedures were used to identify SNPs changing in a parallel or nonparallel way across families. For example, markers that have changed the most in the whole population, markers that have changed differently across families, and genes previously identified as those that have changed in allele frequency. The genomic trajectory taken by each family involves selective sweeps, polygenic changes, hitchhiking, and epistasis. The replicate frequency spectrum was used to measure the similarity of change across families and showed that populations have changed differently. The proportion of markers that reversed direction in allele frequency change varied from 0.00 to 0.02 if the rate of change was greater than 0.02 per generation, or from 0.14 to 0.24 if the rate of change was greater than 0.005 per generation within each family. Cluster-specific SNP effects for stature were estimated using only females and applied to obtain indirect genomic predictions for males. Reranking occurs depending on SNP effects used. Additive genetic correlations between clusters show possible differences in populations. Further research is required to determine how this knowledge can be applied to maintain diversity and optimize selection decisions in the future.

Genomic evaluation with multibreed and crossbred data.

Several types of multibreed genomic evaluation are in use. These include evaluation of crossbreds based on purebred SNP effects, joint evaluation of all purebreds and crossbreds with a single additive effect, and treating each purebred and crossbred group as a separate trait. Additionally, putative quantitative trait nucleotides can be exploited to increase the accuracy of prediction. Existing studies indicate that the prediction of crossbreds based on purebred data has low accuracy, that a joint evaluation can potentially provide accurate evaluations for crossbreds but could lower accuracy for purebreds compared with single-breed evaluations, and that the use of putative quantitative trait nucleotides only marginally increases the accuracy. One hypothesis is that genomic selection is based on estimation of clusters of independent chromosome segments. Subsequently, predicting a particular group type would require a reference population of the same type, and crosses with same breed percentage but different type (F1 vs. F2) would, at best, use separate reference populations. The genomic selection of multibreed population is still an active research topic.

Identifying influential sires and distinct clusters of selection candidates based on genomic relationships to reduce inbreeding in the US Holstein.

High relatedness in the US Holstein breed can be attributed to the increased rate of inbreeding that resulted from strong selection and the extensive use of a few bulls via reproductive biotechnology. The objectives of this study were to determine whether clustering could separate selected candidates into genetically different groups and whether such clustering could reduce the expected inbreeding of the next generation. A genomic relationship matrix composed of 1,145 sires with the most registered progeny in the breed born after 1985 was used for principal component analysis and k-means clustering. The 5 clusters reduced the variance by 25% and contained 171 (C1), 252 (C2), 200 (C3), 244 (C4), and 278 (C5) animals, respectively. The 2 most predominant families were C1 and C2, while C4 contained the most international animals. On average, C1 and C5 contained older animals; the average birth year per cluster was 1988 (C1), 1996 (C2 and C3), 1999 (C4), and 1990 (C5). Increasing to 10 clusters allowed the separation of the predominant sons. Statistically significant differences were observed for indices (net merit index, cheese merit index, and fluid merit index), daughter pregnancy rate, and production traits (milk, fat, and protein), with older clusters having lower merit for production but higher for reproduction. K-means clustering was also used for 20,099 animals considered as selection candidates. Based on the reduction in variance achieved by clustering, 5 to 7 clusters were appropriate. The number of animals in each cluster was 3,577 (C1), 3,073 (C2), 3,302 (C3), 5,931 (C4), and 4,216 (C5). The expected inbreeding from within or across cluster mating was calculated using the complete pedigree, assuming the mean inbreeding of animals born in the same year when parents are unknown. Generally, inbreeding was highest within cluster mating and lowest across cluster mating. Even when 10 clusters were used, one cluster always gave low inbreeding in all scenarios. This suggests that this cluster contains animals that differ from all other groups but still contains enough diversity within itself. Based on lower across cluster inbreeding, up to 7 clusters were appropriate. Statistically significant differences in genomic estimated breeding values were found between clusters. The rankings of clusters for different traits were mostly the same except for reproduction and fat. Results show that diversity within the population exists and clustering of selection candidates can reduce the expected inbreeding of the next generations.

Changes in genetic parameters for milk yield and heat tolerance in the Thai Holstein crossbred dairy population under different heat stress levels and over time.

The objectives of this study were to investigate changes in genetic parameters for milk yield (MY) and heat tolerance of the crossbred Thai Holstein Friesian population under different heat stress levels over time, and to investigate the threshold point of heat stress manifestation on milk production. Genetic parameters were estimated using single-step genomic REML (ssGREML) and traditional REML models. Data included 58,965 test-day MY records from 1999 to 2008 (old data) and 105,485 test-day MY records from 2009 to 2018 (recent data) from the first parity of 24,520 cows. The pedigree included 55,168 animals, of which 882 animals had genotypes. Variance components were estimated with the REMLF90 program using a repeatability model with random regressions on a function of temperature-humidity index (THI) for additive genetic and permanent environmental effects. Fixed effects included farm-calving season combination, breed group-months in milk combination, and age at first calving. Random effects included additive genetic (intercept and slope) effects, permanent environmental (intercept and slope) effects, and herd-month-year of test. The phenotypic mean for MY was 13.33 ± 4.39 kg/d in the old data, and 14.48 ± 4.40 kg/d in the recent data. Estimates over different THI levels for the intercept additive genetic variance using old data ranged from 2.61 to 2.77 and from 5.02 to 5.38 using recent data with the REML method. In ssGREML analyses (performed with recent data only) the estimates for the intercept additive genetic variance ranged from 4.71 to 5.05. Estimates for the slope additive genetic variance were close to zero in all cases, with the largest values (0.024-0.030) at the most extreme THI value (80). Using REML, the covariance between the intercept and the slope additive genetic effects (THI from 72 to 80) ranged from -0.001 to 0.019 with old data and from 0.027 to 0.060 with recent data. The same covariance ranged from 0.026 to 0.057 in ssGREML analyses. The covariance between the intercept and the slope permanent environmental effects ranged from -0.42 to -0.67 for all data and THI levels. Across THI levels, the genetic correlation between MY and heat tolerance varied from -0.06 to 0.13 with old data, from 0.16 to 0.30 with recent data in REML analyses, and from 0.15 to 0.30 in ssGREML analyses, suggesting that in the current population the top animals for MY are more resistant to heat stress. This was expected, because of the introduction of Bos indicus genes in the last years. Heritability estimates for MY ranged from 0.19 to 0.21 (old data) and from 0.33 to 0.40 (recent data) for REML analyses. Heritability estimates for MY using ssGREML ranged from 0.31 to 0.38. A decline in MY was found when the animals' breed composition had more than 97.3% of Holstein genetics, and it was greatest at THI 80. The heritability and genetic correlations observed in this study show that selection for MY is possible without a negative correlated response for heat tolerance. Although the inclusion of genomic information is expected to increase the accuracy of selection, more genotypes must be collected for successful application. Future research should address other production and fitness traits within the Thai Holstein population.

Indirect genomic predictions for milk yield in crossbred Holstein-Jersey dairy cattle.

The objective of this study was to predict genomic breeding values for milk yield of crossbred dairy cattle under different scenarios using single-step genomic BLUP (ssGBLUP). The data set included 13,880,217 milk yield measurements on 6,830,415 cows. Genotypes of 89,558 Holstein, 40,769 Jersey, and 22,373 Holstein-Jersey crossbred animals were used, of which all Holstein, 9,313 Jersey, and 1,667 crossbred animals had phenotypic records. Genotypes were imputed to 45K SNP markers. The SNP effects were estimated from single-breed evaluations for Jersey (JE), Holstein (HO) and crossbreds (CROSS), and multibreed evaluations including all Jersey and Holstein (JE_HO) or approximately equal proportions of Jersey, Holstein, and crossbred animals (MIX). Indirect predictions (IP) of the validation animals (358 crossbred animals with phenotypes excluded from evaluations) were calculated using the resulting SNP effects. Additionally, breed proportions (BP) of crossbred animals were applied as a weight when IP were estimated based on each pure breed. The predictive ability of IP was calculated as the Pearson correlation between IP and phenotypes of the validation animals adjusted for fixed effects in the model. Regression of adjusted phenotypes on IP was used to assess the inflation of IP. The predictive ability of IP for CROSS, JE, HO, JE_HO, and MIX scenario was 0.50, 0.50, 0.47, 0.50, and 0.46, respectively. Using BP was the least successful, with a predictive ability of 0.32. The inflation of the IP for crossbred animals using CROSS, JE, HO, JE_HO, MIX, and BP scenarios were 1.17, 0.65, 0.55, 0.78, 1.00, and 0.85, respectively. The IP of crossbred animals can be predicted using single-step GBLUP under a scenario that includes purebred genotypes.