D.P. Berry

CELL BIOLOGY SYMPOSIUM: Genetics of feed efficiency in dairy and beef cattle1

Increasing food production for the growing human population off a constraining land base will require greater efficiency of production. Genetic improvement of feed efficiency in cattle, which is cumulative and permanent, is one likely vehicle to achieving efficiency gains. The objective of this review is to summarize genetic parameters for feed efficiency traits in dairy and beef cattle and also to address some of the misconceptions associated with feed efficiency in these sectors, as well as discuss the potential use of feed efficiency in breeding programs. A meta-analysis of up to 39 scientific publications in growing cattle clearly showed that genetic variation in feed efficiency exists with a pooled heritability for residual feed intake (RFI) and feed conversion efficiency of 0.33 ± 0.01 (range of 0.07 to 0.62) and 0.23 ± 0.01 (range of 0.06 to 0.46), respectively. Heritability estimates for feed efficiency in cows were lower; a meta-analysis of up to 11 estimates revealed heritability estimates for gross feed efficiency and RFI of 0.06 ± 0.010 and 0.04 ± 0.008, respectively. Meta-analysis of genetic correlations between feed intake, feed efficiency and other performance traits are presented, and selection index theory is used to calculate the proportion of genetic variation in feed intake that can be explained by easy to measure, and often already collected, data. A large proportion of the genetic variation in feed intake could be explained in both growing animals and lactating animals using up to 5 predictor traits, including BW, growth rate, milk yield, body composition, and linear type traits reflecting body size and muscularity. Knowledge of genetic merit for feed intake can be used, along with estimates of genetic merit for energy sinks, to calculate genetic merit for feed efficiency. Therefore, the marginal benefit of collecting actual feed intake data, using the genetic parameters used in this study, appears to be low. There is now sufficient information available to develop a road map on how best to direct research to ensure long-term food security for a growing human population. Gaps in knowledge are identified here, and possibilities to address these gaps are discussed.

Runs of homozygosity and population history in cattle

BackgroundRuns of homozygosity (ROH) are contiguous lengths of homozygous genotypes that are present in an individual due to parents transmitting identical haplotypes to their offspring. The extent and frequency of ROHs may inform on the ancestry of an individual and its population. Here we use high density (n = 777,962) bi-allelic SNPs in a range of cattle breed samples to correlate ROH with the pedigree-based inbreeding coefficients and to validate subsequent analyses using 54,001 SNP genotypes. This study provides a first testing of the inference drawn from ROH through comparison with estimates of inbreeding from calculations based on the detailed pedigree data available for several breeds.ResultsAll animals genotyped on the HD panel displayed at least one ROH that was between 1–5 Mb in length with certain regions of the genome more likely to be involved in a ROH than others. Strong correlations (r = 0.75, p < 0.0001) existed between the pedigree-based inbreeding coefficient and a statistic based on sum of ROH of length > 0.5 KB and suggests that in the absence of an animal’s pedigree data, the extent of a genome under ROH may be used to infer aspects of recent population history even from relatively few samples.ConclusionsOur findings suggest that ROH are frequent across all breeds but differing patterns of ROH length and burden illustrate variations in breed origins and recent management.

Mid-infrared prediction of bovine milk fatty acids across multiple breeds, production systems, and countries

Increasing consumer concern exists over the relationship between food composition and human health. Because of the known effects of fatty acids on human health, the development of a quick, inexpensive, and accurate method to directly quantify the fatty acid (FA) composition in milk would be valuable for milk processors to develop a payment system for milk pertinent to their customer requirements and for farmers to adapt their feeding systems and breeding strategies accordingly. The aim of this study was (1) to confirm the ability of mid-infrared spectrometry (MIR) to quantify individual FA content in milk by using an innovative procedure of sampling (i.e., samples were collected from cows belonging to different breeds, different countries, and in different production systems); (2) to compare 6 mathematical methods to develop robust calibration equations for predicting the contents of individual FA in milk; and (3) to test interest in using the FA equations developed in milk as basis to predict FA content in fat without corrections for the slope and the bias of the developed equations. In total, 517 samples selected based on their spectral variability in 3 countries (Belgium, Ireland, and United Kingdom) from various breeds, cows, and production systems were analyzed by gas chromatography (GC). The samples presenting the largest spectral variability were used to calibrate the prediction of FA by MIR. The remaining samples were used to externally validate the 28 FA equations developed. The 6 methods were (1) partial least squares regression (PLS); (2) PLS+repeatability file (REP); (3) first derivative of spectral data+PLS; (4) first derivative+REP+PLS; (5) second derivative of spectral data+PLS; and (6) second derivative+REP+PLS. Methods were compared on the basis of the cross-validation coefficient of determination (R2cv), the ratio of standard deviation of GC values to the standard error of cross-validation (RPD), and the validation coefficient of determination (R2v). The third and fourth methods had, on average, the highest R2cv, RPD, and R2v. The final equations were built using all GC and the best accuracy was observed for the infrared predictions of C4:0, C6:0, C8:0, C10:0, C12:0, C14:0, C16:0, C18:0, C18:1 trans, C18:1 cis-9, C18:1 cis, and for some groups of FA studied in milk (saturated, monounsaturated, unsaturated, short-chain, medium-chain, and long-chain FA). These equations showed R2cv greater than 0.95. With R2cv equal to 0.85, the MIR prediction of polyunsaturated FA could be used to screen the cow population. As previously published, infrared predictions of FA in fat are less accurate than those developed from FA content in milk (g/dL of milk) and no better results were obtained by using milk FA predictions if no corrections for bias and slope based on reference milk samples with known contents of FA were used. These results indicate the usefulness of equations with R2cv greater than 95% in milk payment systems and the usefulness of equations with R2cv greater than 75% for animal breeding purposes.

Phenotypic and genetic parameters for different measures of feed efficiency in different breeds of Irish performance-tested beef bulls

No genetic parameters for performance and feed efficiency traits are available for Irish performance-tested bulls. The objective of this study was to determine the phenotypic and genetic variation for feed intake, BW, ADG, and measures of feed efficiency including feed conversion ratio (FCR), relative growth rate, Kleiber ratio, residual BW gain (RG), and residual feed intake (RFI). Observations were available on up to 2,605 bulls for each trait from one test station across 24 yr; breeds included in the analyses were Aberdeen Angus (AN), Charolais (CH), Hereford, Limousin (LI), and Simmental. The test period was at least 70 d. Bulls were individually offered concentrates ad libitum, with a restricted forage allowance. Differences in performance and feed efficiency existed among breeds. For example, AN, on average, ate 0.04 kg of DM/d more than CH but had ADG of 0.14 kg/d less over the 70-d test period. Results showed LI and CH were the most efficient breeds when efficiency was defined as FCR or RFI. When animals were partitioned into groups based on high, medium, or low RFI, the low RFI (i.e., most efficient) group were also the more efficient as defined by RG and FCR. The low RFI group had the same ADG as the medium group and a greater ADG (P < 0.01) than the high group (1.67 vs. 1.66 and 1.63 kg/d); yet they ate 0.67 kg of DM/d less (P < 0.001) than the medium RFI group and 1.22 kg of DM/d less (P < 0.001) than the high RFI (i.e., least efficient) group. Genetic parameters for all performance and efficiency measures were estimated across breeds using linear animal mixed models; heritability estimates for feed efficiency traits ranged from 0.28 +/- 0.06 (RG) to 0.45 +/- 0.06 (RFI). An additional series of analyses included a maternal component in the model; maternal heritability estimates for feed efficiency traits ranged from 0.05 +/- 0.03 (RG) to 0.11 +/- 0.05 (relative growth rate). Genetic correlations between most of the different feed efficiency measures were strong. Results from this study indicate significant genetic differences in performance and some measures of feed efficiency among performance-tested beef bulls.

Low numbers of ovarian follicles ≥3 mm in diameter are associated with low fertility in dairy cows

The total number of ovarian follicles ≥ 3mm in diameter (antral follicle count, AFC) during follicular waves varies among cattle of similar age, but AFC is highly repeatable within individuals. We hypothesized that lower AFC could be associated with reduced fertility in cattle. The AFC was assessed by ultrasonography for 2 d consecutively during the first wave of follicular growth of the estrous cycle, 4.6±1.43 d (mean ± SD) after estrus, in 306 Holstein-Friesian dairy cows approximately 70 d postpartum. Cows were classified into 3 groups based on AFC: low (AFC ≤15), intermediate (AFC=16 to 24), and high (AFC ≥25). During the cycle in which AFC was assessed and in subsequent cycles, cows were artificially inseminated (AI) following detection of estrus, and pregnancy status was assessed using ultrasonography. Cows with high AFC had 3.34 times greater odds of being pregnant at the end of the breeding season compared with cows with low AFC; the odds of a successful pregnancy at first service were 1.75 times greater in the intermediate compared with the low group. The predicted probability of a successful pregnancy by the end of the breeding period (length of breeding season was 86±16.3 d) was 94, 88, and 84% for the high, intermediate, and low AFC groups, respectively. No difference was evident among groups in 21-d submission rate (proportion of all cows detected in estrus and submitted for AI in the first 21 d of the breeding season), but the interval from calving to conception was shorter in the high (109.5±5.1 d) versus low (117.1±4 d) group, and animals with intermediate AFC received fewer services during the breeding season (2.3±0.1) compared with animals with low AFC (2.7±0.1). Lactating cows with ≤15 ovarian follicles have lower reproductive performance compared with cows with higher numbers of follicles, but the existence of a positive association between high numbers of ovarian follicles and fertility is yet to be established.

Genetics and genomics of reproductive performance in dairy and beef cattle

Excellent reproductive performance in both males and females is fundamental to profitable dairy and beef production systems. In this review we undertook a meta-analysis of genetic parameters for female reproductive performance across 55 dairy studies or populations and 12 beef studies or populations as well as across 28 different studies or populations for male reproductive performance. A plethora of reproductive phenotypes exist in dairy and beef cattle and a meta-analysis of the literature suggests that most of the female reproductive traits in dairy and beef cattle tend to be lowly heritable (0.02 to 0.04). Reproductive-related phenotypes in male animals (e.g. semen quality) tend to be more heritable than female reproductive phenotypes with mean heritability estimates of between 0.05 and 0.22 for semen-related traits with the exception of scrotal circumference (0.42) and field non-return rate (0.001). The low heritability of reproductive traits, in females in particular, does not however imply that genetic selection cannot alter phenotypic performance as evidenced by the decline until recently in dairy cow reproductive performance attributable in part to aggressive selection for increased milk production. Moreover, the antagonistic genetic correlations among reproductive traits and both milk (dairy cattle) and meat (beef cattle) yield is not unity thereby implying that simultaneous genetic selection for both increased (milk and meat) yield and reproductive performance is indeed possible. The required emphasis on reproductive traits within a breeding goal to halt deterioration will vary based on the underlying assumptions and is discussed using examples for Ireland, the United Kingdom and Australia as well as quantifying the impact on genetic gain for milk production. Advancements in genomic technologies can aid in increasing the accuracy of selection for especially reproductive traits and thus genetic gain. Elucidation of the underlying genomic mechanisms for reproduction could also aid in resolving genetic antagonisms. Past breeding programmes have contributed to the deterioration in reproductive performance of dairy and beef cattle. The tools now exist, however, to reverse the genetic trends in reproductive performance underlying the observed phenotypic trends.

Residual intake and body weight gain: A new measure of efficiency in growing cattle

Interest in improving feed efficiency in cattle is intensifying. Residual feed intake (RFI), which is the difference between expected intake and that predicted based on energy demands, is now the most commonly used measure of feed efficiency over a given time period. However, RFI, as commonly defined, is independent of growth rate, which may affect its acceptance by industry. Residual BW gain (RG) has also been proposed as a measure of feed efficiency and is represented as the residuals from a multiple regression model regressing ADG on both DMI and BW. In this study, we propose a new trait, residual intake and BW gain (RIG), which retains the favorable characteristic of both RFI and RG being independent of BW, but animals superior for RIG have, on average, both greater ADG and reduced DMI. Phenotypic and genetic analyses were undertaken on up to 2,605 purebred performance-tested bulls. Clear phenotypic differences in DMI and ADG existed between animals divergent for RIG. The heritability of RIG was 0.36 ± 0.06, which is consistent with the heritability estimates of RFI and other feed efficiency traits measured in the study. The RIG trait was both phenotypically and genetically negatively correlated with DMI and positively correlated with ADG; no correlation existed between RIG and BW. The advantages of both reduced daily DMI and greater ADG in animals superior for RIG are demonstrated compared with animals superior for either RFI or RG.

Genome-wide associations for milk production and somatic cell score in Holstein-Friesian cattle in Ireland

BackgroundContemporary dairy breeding goals have broadened to include, along with milk production traits, a number of non-production-related traits in an effort to improve the overall functionality of the dairy cow. Increased indirect selection for resistance to mastitis, one of the most important production-related diseases in the dairy sector, via selection for reduced somatic cell count has been part of these broadened goals. A number of genome-wide association studies have identified genetic variants associated with milk production traits and mastitis resistance, however the majority of these studies have been based on animals which were predominantly kept in confinement and fed a concentrate-based diet (i.e. high-input production systems). This genome-wide association study aims to detect associations using genotypic and phenotypic data from Irish Holstein-Friesian cattle fed predominantly grazed grass in a pasture-based production system (low-input).ResultsSignificant associations were detected for milk yield, fat yield, protein yield, fat percentage, protein percentage and somatic cell score using separate single-locus, frequentist and multi-locus, Bayesian approaches. These associations were detected using two separate populations of Holstein-Friesian sires and cows. In total, 1,529 and 37 associations were detected in the sires using a single SNP regression and a Bayesian method, respectively. There were 103 associations in common between the sires and cows across all the traits. As well as detecting associations within known QTL regions, a number of novel associations were detected; the most notable of these was a region of chromosome 13 associated with milk yield in the population of Holstein-Friesian sires.ConclusionsA total of 276 of novel SNPs were detected in the sires using a single SNP regression approach. Although obvious candidate genes may not be initially forthcoming, this study provides a preliminary framework upon which to identify the causal mechanisms underlying the various milk production traits and somatic cell score. Consequently this will deepen our understanding of how these traits are expressed.

Prevalence of, and risk factors associated with, perinatal calf mortality in pasture-based Holstein-Friesian cows

Recent publications indicate that the prevalence of perinatal mortality has increased in some dairy industries and an increased proportion of this loss is not associated with the traditional risk factors for perinatal mortality. The objectives of this study were to establish the prevalence of perinatal mortality (calf death within 24 h of calving) in Irish dairy herds and to determine the current significance of putative risk factors in pasture-based management systems. A total of 182 026 records of full-term calvings from Holstein-Friesian dams served by artificial insemination (AI) sires of seven breeds in herds of 20 calvings or more per year were available from the Irish national breeding database over 4 years (2002 to 2005). The prevalence of perinatal mortality was 4.29% (7.7% in primiparae and 3.5% in pluriparae). The likelihood of perinatal mortality increased between 2002 and 2005 and was greatest in June and in winter. There was an interaction (P < 0.001) between the effect of calving assistance and parity with the effect of dystocia on perinatal mortality being greater in primiparae. The odds of perinatal mortality were greater in male (OR = 1.12; P < 0.001) and in twin calves (OR = 5.70-13.36; P < 0.001) and in dams that had perinatal mortality at the previous calving (OR = 4.21; P < 0.001). The logit of the probability of perinatal mortality increased by 0.099 per unit increase in sire predicted transmitting ability (PTA) for direct perinatal mortality. The probability of perinatal mortality increased at an increasing rate in primiparae as animals calved at a younger age relative to the median age at first calving. The only herd-level factor examined, herd size did not affect the odds of perinatal mortality. These data indicate that the prevalence of perinatal mortality in this cattle population is similar to that in other pasture-based dairy systems worldwide. The putative exposures and attributes traditionally associated with perinatal mortality were associated with perinatal mortality in this pasture-based dairy cow population. The practical implication of these results is that as many of the significant risk factors are largely not under management control (year of calving, month of calving, twin calving, primiparity, previous perinatal mortality and foetal gender), herd owners must focus on the significant determinants under their control (age at first calving, sire genetic merit for direct perinatal mortality and both the extent of calving supervision and the degree of assistance), in order to reduce the prevalence of perinatal mortality and improve perinatal welfare.

Genetics of tuberculosis in Irish Holstein-Friesian dairy herds

Information is lacking on genetic parameters for tuberculosis (TB) susceptibility in dairy cattle. Mycobacterium bovis is the principal agent of tuberculosis in cattle. The objective of this study was to quantify the genetic variation present among Irish Holstein-Friesian dairy herds in their susceptibility to M. bovis infection. A total of 15,182 cow and 8,104 heifer single intradermal comparative tuberculin test (SICTT, a test for M. bovis exposure and presumed infection) records from November 1, 2002, to October 31, 2005, were available for inclusion in the analysis. Data on observed carcass TB lesions from abattoirs were also available for inclusion in the analysis. The only animals retained were those present in a herd during episodes in which at least 2 animals showed evidence of infection; this ensured a high likelihood of exposure to M. bovis. Linear animal models, and sire and animal threshold models were used to estimate the variance components for susceptibility to M. bovis-purified protein derivative (PPD) responsiveness and confirmed M. bovis infection. The heritability estimates from the threshold sire models were biased upward because the relatedness between dam-daughter pairs was ignored. The threshold animal model produced heritability estimates of 0.14 in cows and 0.12 in heifers for susceptibility to M. bovis-PPD responsiveness, and 0.18 in cows for confirmed M. bovis infection susceptibility. Therefore, exploitable genetic variation exists among Irish dairy cows for susceptibility to M. bovis infection. Sire rankings from the linear and threshold animal models were similar, indicating that either model could be used for the analysis of susceptibility to M. bovis-PPD responsiveness. A favorable genetic correlation close to unity was observed between susceptibility to confirmed M. bovis infection and M. bovis-PPD responsiveness, indicating that direct selection for resistance to M. bovis-PPD responsiveness will indirectly reduce susceptibility to confirmed M. bovis infection. Data from the national TB eradication program could be used routinely to estimate breeding values for susceptibility to M. bovis infection.