J.E. Pryce

A validated genome-wide association study in 2 dairy cattle breeds for milk production and fertility traits using variable length haplotypes

Genome-wide association studies (GWAS) were used to discover genomic regions explaining variation in dairy production and fertility traits. Associations were detected with either single nucleotide polymorphism (SNP) markers or haplotypes of SNP alleles. An across-breed validation strategy was used to narrow the genomic interval containing causative mutations. There were 39,048 SNP tested in a discovery population of 780 Holstein sires and validated in 386 Holsteins and 364 Jersey sires. Previously identified mutations affecting milk production traits were confirmed. In addition, several novel regions were identified, including a putative quantitative trait loci for fertility on chromosome 18 that was detected only using haplotypes greater than 3 SNP long. It was found that the precision of quantitative trait loci mapping increased with haplotype length as did the number of validated haplotypes discovered, especially across breed. Promising candidate genes have been identified in several of the validated regions.

Multivariate analysis of a genome-wide association study in dairy cattle.

Multiple-trait genome-wide association study (GWAS) analyses were compared with single-trait GWAS for power to discover and subsequently validate genetic markers (single nucleotide polymorphisms; SNP) associated with dairy traits. The SNP associations were discovered in 1 Holstein population and validated in both a Holstein population consisting of bulls younger than those in the discovery population and a Jersey population. The multivariate methods used were a principal component analysis and a series of bivariate analyses. The statistical power of detecting associations using multiple-trait GWAS was as good as or better than that of the best single-trait GWAS. Additional SNP associations were found with the multivariate methods that had not been discovered in the single-trait analyses; this was achieved without an increase in the false discovery rate. From the multivariate analysis, 4 common pleiotropic patterns were identified among the putative quantitative trait loci (QTL) affecting the Australian selection index. These patterns could be interpreted as a primary effect of the putative QTL on 1 or more milk components and secondary effects on other components. The multivariate analysis did not appear to increase the precision with which putative QTL were mapped.

Variation in residual feed intake in Holstein-Friesian dairy heifers in southern Australia

Feed conversion efficiency of dairy cattle is an important component of the profitability of dairying, given that the cost of feed accounts for much of total farm expenses. Residual feed intake (RFI) is a useful measure of feed conversion efficiency, as it can be used to compare individuals with the same or differing levels of production during the period of measurement. If genetic variation exists in RFI among dairy cattle, selection for lower RFI could improve profitability. In this experiment, RFI was defined as the difference between an animals actual feed intake and its expected feed intake, which was determined by regression of dry matter (DM) intake against mean body weight (BW) and growth rate. Nine hundred and three Holstein-Friesian heifer calves, aged between 5 and 7 mo, were measured for RFI in 3 cohorts of approximately 300 animals. Calves were housed under feedlot style conditions in groups of 15 to 20 for 85 to 95 d and had ad libitum access to a cubed alfalfa hay. Intakes of individual animals were recorded via an electronic feed recording system and BW gain was determined by weighing animals once or twice weekly, over a period of 60 to 70 d. Calves had DM intake (mean ± SD) of 8.3±1.37 kg of DM/d over the measurement period with BW gains of 1.1±0.17 kg/d. In terms of converting feed energy for maintenance and growth, the 10% most efficient calves (lowest RFI) ate 1.7 kg of DM less each day than the 10% least efficient calves (highest RFI) for the same rate of growth. Low-RFI heifers also had a significantly lower rate of intake (g/min) than high-RFI heifers. The heritability estimate of RFI (mean ± SE) was 0.27 (±0.12). These results indicate that substantial genetic variation in RFI exists, and that the magnitude of this variation is large enough to enable this trait to be considered as a candidate trait for future dairy breeding goals. A primary focus of future research should be to ensure that calves that are efficient at converting feed energy for maintenance and growth also become efficient at converting feed energy to milk. Future research will also be necessary to identify the consequences of selection for RFI on other traits (especially fertility and other fitness traits) and if any interactions exist between RFI and feeding level.

Short communication: Genomic selection using a multi-breed, across-country reference population

Three breeds (Fleckvieh, Holstein, and Jersey) were included in a reference population, separately and together, to assess the accuracy of prediction of genomic breeding values in single-breed validation populations. The accuracy of genomic selection was defined as the correlation between estimated breeding values, calculated using phenotypic data, and genomic breeding values. The Holstein and Jersey populations were from Australia, whereas the Fleckvieh population (dual-purpose Simmental) was from Austria and Germany. Both a BLUP with a multi-breed genomic relationship matrix (GBLUP) and a Bayesian method (BayesA) were used to derive the prediction equations. The hypothesis tested was that having a multi-breed reference population increased the accuracy of genomic selection. Minimal advantage existed of either GBLUP or BayesA multi-breed genomic evaluations over single-breed evaluations. However, when the goal was to predict genomic breeding values for a breed with no individuals in the reference population, using 2 other breeds in the reference was generally better than only 1 breed.

Invited review: overview of new traits and phenotyping strategies in dairy cattle with a focus on functional traits

For several decades, breeding goals in dairy cattle focussed on increased milk production. However, many functional traits have negative genetic correlations with milk yield, and reductions in genetic merit for health and fitness have been observed. Herd management has been challenged to compensate for these effects and to balance fertility, udder health and metabolic diseases against increased production to maximize profit without compromising welfare. Functional traits, such as direct information on cow health, have also become more important because of growing concern about animal well-being and consumer demands for healthy and natural products. There are major concerns about the impact of drugs used in veterinary medicine on the spread of antibiotic-resistant strains of bacteria that can negatively impact human health. Sustainability and efficiency are also increasingly important because of the growing competition for high-quality, plant-based sources of energy and protein. Disruptions to global environments because of climate change may encourage yet more emphasis on these traits. To be successful, it is vital that there be a balance between the effort required for data recording and subsequent benefits. The motivation of farmers and other stakeholders involved in documentation and recording is essential to ensure good data quality. To keep labour costs reasonable, existing data sources should be used as much as possible. Examples include the use of milk composition data to provide additional information about the metabolic status or energy balance of the animals. Recent advances in the use of mid-infrared spectroscopy to measure milk have shown considerable promise, and may provide cost-effective alternative phenotypes for difficult or expensive-to-measure traits, such as feed efficiency. There are other valuable data sources in countries that have compulsory documentation of veterinary treatments and drug use. Additional sources of data outside of the farm include, for example, slaughter houses (meat composition and quality) and veterinary labs (specific pathogens, viral loads). At the farm level, many data are available from automated and semi-automated milking and management systems. Electronic devices measuring physiological status or activity parameters can be used to predict events such as oestrus, and also behavioural traits. Challenges concerning the predictive biology of indicator traits or standardization need to be solved. To develop effective selection programmes for new traits, the development of large databases is necessary so that high-reliability breeding values can be estimated. For expensive-to-record traits, extensive phenotyping in combination with genotyping of females is a possibility.

Deterministic models of breeding scheme designs that incorporate genomic selection

A deterministic model to calculate rates of genetic gain and inbreeding was used to compare a range of breeding scheme designs under genomic selection (GS) for a population of 140,000 cows. For most schemes it was assumed that the reliability of genomic breeding values (GEBV) was 0.6 across 4 pathways of selection. In addition, the effect of varying reliability on the ranking of schemes was also investigated. The schemes considered included intense selection in male pathways and genotyping of 1,000 young bulls (GS-Y). This scheme was extended to include selection in females and to include a worldwide scheme similar to GS-Y, but 6 times as large and assuming genotypes were freely exchanged between 6 countries. An additional worldwide scheme was modeled where GEBV were available through international genetic evaluations without exchange of genotypes. Finally, a closed nucleus herd that used juvenile in vitro embryo transfer in heifers was modeled so that the generation interval in female pathways was reduced to 1 or 2 yr. When the breeding schemes were compared using a GEBV reliability of 0.6, the rates of genetic gain were between 59 and 130% greater than the rate of genetic gain achieved in progeny testing. This was mainly through reducing the generation interval and increasing selection intensity. Genomic selection of females resulted in a 50% higher rate of genetic gain compared with restricting GS to young bulls only. The annual rates of inbreeding were, in general, 60% lower than with progeny testing, because more sires of bulls and sires of cows were selected, thus increasing the effective population size. The exception was in nucleus breeding schemes that had very short generation intervals, resulting in higher rates of both gain and inbreeding. It is likely that breeding companies will move rapidly to alter their breeding schemes to make use of genomic selection because benefits to the breeding companies and to the industry are considerable.

Analyzing the heterogeneity of farmers' preferences for improvements in dairy cow traits using farmer typologies.

Giving consideration to farmers preferences for improvements in animal traits when designing genetic selection tools such as selection indexes might increase the uptake of these tools. The increase in use of genetic selection tools will, in turn, assist in the realization of genetic gain in breeding programs. However, the determination of farmers preferences is not trivial because of its large heterogeneity. The aim of this study was to quantify Australian dairy farmers preferences for cow trait improvements to inform and ultimately direct the choice of traits and selection indexes in the 2014 review of the National Breeding Objective. A specific aim was to analyze the heterogeneity of preferences for cow trait improvements by determining whether there are farmer types that can be identified with specific patterns of preferences. We analyzed whether farmer types differed in farming system, socioeconomic profile, and attitudes toward breeding and genetic evaluation tools. An online survey was developed to explore farmers preferences for improvement in 13 cow traits. The pairwise comparisons method was used to derive a ranking of the traits for each respondent. A total of 551 farmers fully completed the survey. A principal component analysis followed by a Ward hierarchical cluster analysis was used to group farmers according to their preferences. Three types of farmers were determined: (1) production-focused farmers, who gave the highest preference of all for improvements in protein yield, lactation persistency, feed efficiency, cow live weight, and milking speed; (2) functionality-focused farmers with the highest preferences of all for improvements in mastitis, lameness, and calving difficulty; and (3) type-focused farmers with the highest preferences of all for mammary system and type. Farmer types differed in their age, their attitudes toward genetic selection, and in the selection criteria they use. Surprisingly, farmer types did not differ for herd size, calving, feeding system, or breed. These results support the idea that preferences for cow trait improvements are intrinsic to farmers and not to production systems or breeds. As a result of this study, and some bioeconomic modeling (not included in this study), the Australian dairy industry has implemented a main index and 2 alternative indexes targeting the different farmer types described here.

New breeding objectives and selection indices for the Australian dairy industry

This study comprises an update of the economic values for dairy traits for the Australian industry and the formulation of updated selection indices. An economic model, which calculates partial economic values for each trait individually, was developed to determine the economic implications of selective dairy breeding, based on the effect of trait changes on the profit of commercial dairy farms in Australia. Selection indices were developed from economic values, which were transformed into base economic weights by including the discounted genetic expressions coefficients. Economic weights (in Australian dollars) were 1.79, 6.92, -0.10, -5.44, 8.84, 7.68, 1.07, 4.86, 1.91, 3.51, 4.90, 0.31, 2.03, 2.00, and 0.59, for milk fat (kg), milk protein (kg), milk volume (L), body weight (kg), survival (%), residual survival (%), somatic cell count (cells/mL), fertility (%), mammary system [Australian Breeding Value (ABV) unit], temperament (ABV unit), milking speed (ABV unit), udder depth (%), overall type (%), fore udder attachment (%), and pin set (%), respectively. The updated economic weights presented in this study constituted the basis of the definition for 3 new indices. These indices were developed from combination of bioeconomic principles, patterns of farmer preferences for trait improvements, and desired gains approaches. The 3 indices, Balanced Performance Index, Health Weighted Index, and Type Weighted Index, have been released to the industry.

Genetic markers for lactation persistency in primiparous Australian dairy cows.

Good performance in extended lactations of dairy cattle may have a beneficial effect on food costs, health, and fertility. Because data for extended lactation performance is scarce, lactation persistency has been suggested as a suitable selection criterion. Persistency phenotypes were calculated in several ways: P1 was yield relative to an approximate peak, P2 was the slope after peak production, and P3 was a measure derived to be phenotypically uncorrelated to yield and calculated as a function of linear regressions on test-day deviations of days in milk. Phenotypes P1, P2, and P3 were calculated for sires as solutions estimated from a random regression model fitted to milk yield. Because total milk yield, calculated as the sum of daily sire solutions, was correlated to P1 and P2 (r=0.30 and 0.35 for P1 and P2, respectively), P1 and P2 were also adjusted for milk yield (P1adj and P2adj, respectively). To find genomic regions associated with the persistency phenotypes, we used a discovery population of 743 Holstein bulls proven before 2005 and 2 validation data sets of 357 Holstein bulls proven after 2005 and 294 Jersey sires. Two strategies were used to search for genomic regions associated with persistency: 1) persistency solutions were regressed on each single nucleotide polymorphism (SNP) in turn and 2) a genomic selection method (BayesA) was used where all SNP were fitted simultaneously. False discovery rates in the validation data were high (>66% in Holsteins and >77% in Jerseys). However, there were 2 genomic regions on chromosome 6 that validated in both breeds, including a cluster of 6 SNP at around 13.5 to 23.7 Mbp and another cluster of 5 SNP (70.4 to 75.6 Mbp). A third cluster validated in both breeds on chromosome 26 (0.33 to 1.46 Mbp). Validating SNP effects across 2 breeds is unlikely to happen by chance even when false discovery rates within each breed are high. However, marker-assisted selection on these selected SNP may not be the best way to improve this trait because the average variation explained by validated SNP was only 1 to 2%. Genomic selection could be a better alternative. Correlations between genomic breeding values predicted using all SNP simultaneously and estimated breeding values based on progeny test were twice as high as the equivalent correlations between estimated breeding values and parent average. Persistency is a good candidate for genomic selection because the trait is expressed late in lactation.

Prediction of liveweight of cows from type traits and its relationship with production and fitness traits

Liveweight (LWT) data for Australian Holstein cows was predicted from different type traits based on actual LWT and type data of 932 cows collected from 20 different herds over a 3-yr period. In addition to LWT measured using scales, visual estimates of LWT were also available on 90% of the cows with LWT data. The future predictive ability of different models was assessed using 10-fold cross-validation. The relationships between LWT and selected type traits, including body condition score (BCS), were also calculated to assess the usefulness of various traits to predict LWT genetically. The relationships of predicted LWT measures with production and fitness traits were also estimated in an attempt to assess the consequence of continuous selection on an economic index that includes predicted LWT with a negative economic value. The heritability of LWT was 0.4. Among type traits, stature, chest width, bone quality, BCS, udder depth, central ligament, and muzzle width were correlated with LWT both genetically and phenotypically and were used to predict LWT of cows. Predicted LWT measures, using several sets of traits and visually estimated LWT were genetically highly correlated with each other (>0.73). Phenotypically, visually estimated LWT of cows was slightly more correlated with actual LWT than that predicted from type traits, but genetically both approaches gave the same accuracy. The predicted estimates of LWT were also positively genetically correlated with energy-corrected milk yield and had near zero correlation with survival. The correlations of different measures of LWT with fertility traits were unfavorable or near zero, suggesting that selection for reduced LWT may not cause deterioration in fertility traits. However, it may be useful to consider broadening the breeding objective to include traits that are associated with energy balance, particularly if traits such as BCS and bone quality are included in the set of traits used to predict LWT. Based on the results from this study, the inclusion of predicted LWT with negative economic values into the breeding objective would have no negative effect on fitness traits.