The relationships between early lactation energy status indicators and endocrine fertility traits in dairy cows.

The relationships between dairy cow milk-based energy status (ES) indicators and fertility traits were studied during periods 8 to 21, 22 to 35, 36 to 49, and 50 to 63 d in milk. Commencement of luteal activity (C-LA) and interval from calving to the first heat (CFH), based on frequent measurements of progesterone by the management tool Herd Navigator (DeLaval), were used as fertility traits. Energy status indicator traits were milk ß-hydroxybutyrate (BHB) concentration provided by Herd Navigator and milk fat:protein ratio, concentration of C18:1 cis-9, the ratio of fatty acids (FA) C18:1 cis-9 and C10:0 in test-day milk samples, and predicted plasma concentration of nonesterified fatty acids (NEFA) on test days. Plasma NEFA predictions were based either directly on milk mid-infrared spectra (MIR) or on milk fatty acids based on MIR spectra (NEFAmir and NEFAfa, respectively). The average (standard deviation) C-LA was 39.3 (±16.6) days, and the average CFH was 50.7 (±17.2) days. The correlations between fertility traits and ES indicators tended to be higher for multiparous (r < 0.28) than for primiparous (r < 0.16) cows. All correlations were lower in the last period than in the other periods. In period 1, correlations of C-LA with NEFAfa and BHB, respectively, were 0.15 and 0.14 for primiparous and 0.26 and 0.22 for multiparous cows. The associations between fertility traits and ES indicators indicated that negative ES during the first weeks postpartum may delay the onset of luteal activity. Milk FPR was not as good an indicator for cow ES as other indicators. According to these findings, predictions of plasma NEFA and milk FA based on milk MIR spectra of routine test-day samples and the frequent measurement of milk BHB by Herd Navigator gave equally good predictions of cow ES during the first weeks of lactation. Our results indicate that routinely measured milk traits can be used for ES evaluation in early lactation.

Reliability of breeding values for feed intake and feed efficiency traits in dairy cattle: When dry matter intake recordings are sparse under different scenarios.

Currently, routine recordings of dry matter intake (DMI) in commercial herds are practically nonexistent. Recording DMI from commercial herds is a prerequisite for the inclusion of feed efficiency (FE) traits in dairy cattle breeding goals. To develop future on-farm phenotyping strategies, recording strategies that are low cost and less demanding logistically and that give relatively accurate estimates of the animal's genetic merit are therefore needed. The objectives of this study were (1) to estimate genetic parameters for daily DMI and FE traits and use the estimated parameters to simulate daily DMI phenotypes under different DMI recording scenarios (SCN) and (2) to use the simulated data to estimate for different scenarios the associated reliability of estimated breeding value and accuracies of genomic prediction for varying sizes of reference populations. Five on-farm daily DMI recording scenarios were simulated: once weekly (SCN1), once monthly (SCN2), every 2 mo (SCN3), every 3 mo (SCN4), and every 4 mo (SCN5). To estimate reliability of estimated breeding values, DMI and FE observations and true breeding values were simulated based on variance components estimated for daily observations of Nordic Red cows. To emulate realistic on-farm recording, 5 data set replicates, each with 36,037 DMI and FE records, were simulated for real pedigree and data structure of 789 Holstein cows. Observations for the 5 DMI recording scenarios were generated by discarding data in a step-wise manner from the full simulated data per the scenario's definitions. For each of these scenarios, reliabilities were calculated as correlation between the true and estimated breeding values. Variance components and genetic parameters were estimated for daily DMI, residual feed intake (RFI), and energy conversion efficiency (ECE) fitting the random regression model. Data for variance components were from 227 primiparous Nordic Red dairy cows covering 8 to 280 d in milk. Lactation-wise heritability for DMI, RFI, and ECE was 0.33, 0.12, and 0.32, respectively, and daily heritability estimates during lactation ranged from 0.18 to 0.45, 0.08 to 0.32, and 0.08 to 0.45 for DMI, RFI, and ECE, respectively. Genetic correlations for DMI between different stages of lactation ranged from -0.50 to 0.99. The comparison of different on-farm DMI recording scenarios indicated that adopting a less-frequent recording scenario (SCN3) gave a similar level of accuracy as SCN1 when 17 more daughters are recorded per sire over the 46 needed for SCN1. Such a strategy is less demanding logistically and is low cost because fewer observations need to be collected per animal. The accuracy of genomic predictions associated with the 5 recording scenarios indicated that setting up a relatively larger reference population and adopting a less-frequent DMI sampling scenario (e.g., SCN3) is promising. When the same reference population size was considered, the genomic prediction accuracy of SCN3 was only 5.0 to 7.0 percentage points lower than that for the most expensive DMI recording strategy (SCN1). We concluded that DMI recording strategies that are sparse in terms of records per cow but with slightly more cows recorded per sire are advantageous both in genomic selection and in traditional progeny testing schemes when accuracy, logistics, and cost implications are considered.

Body and milk traits as indicators of dairy cow energy status in early lactation.

The inclusion of feed intake and efficiency traits in dairy cow breeding goals can lead to increased risk of metabolic stress. An easy and inexpensive way to monitor postpartum energy status (ES) of cows is therefore needed. Cows' ES can be estimated by calculating the energy balance from energy intake and output and predicted by indicator traits such as change in body weight (ΔBW), change in body condition score (ΔBCS), milk fat:protein ratio (FPR), or milk fatty acid (FA) composition. In this study, we used blood plasma nonesterified fatty acids (NEFA) concentration as a biomarker for ES. We determined associations between NEFA concentration and ES indicators and evaluated the usefulness of body and milk traits alone, or together, in predicting ES of the cow. Data were collected from 2 research herds during 2013 to 2016 and included 137 Nordic Red dairy cows, all of which had a first lactation and 59 of which also had a second lactation. The data included daily body weight, milk yield, and feed intake and monthly BCS. Plasma samples for NEFA were collected twice in lactation wk 2 and 3 and once in wk 20. Milk samples for analysis of fat, protein, lactose, and FA concentrations were taken on the blood sampling days. Plasma NEFA concentration was higher in lactation wk 2 and 3 than in wk 20 (0.56 ± 0.30, 0.43 ± 0.22, and 0.13 ± 0.06 mmol/L, respectively; all means ± standard deviation). Among individual indicators, C18:1 cis-9 and the sum of C18:1 in milk had the highest correlations (r = 0.73) with NEFA. Seven multiple linear regression models for NEFA prediction were developed using stepwise selection. Of the models that included milk traits (other than milk FA) as well as body traits, the best fit was achieved by a model with milk yield, FPR, ΔBW, ΔBCS, FPR × ΔBW, and days in milk. The model resulted in a cross-validation coefficient of determination (R2cv) of 0.51 and a root mean squared error (RMSE) of 0.196 mmol/L. When only milk FA concentrations were considered in the model, NEFA prediction was more accurate using measurements from evening milk than from morning milk (R2cv = 0.61 vs. 0.53). The best model with milk traits contained FPR, C10:0, C14:0, C18:1 cis-9, C18:1 cis-9 × C14:0, and days in milk (R2cv = 0.62; RMSE = 0.177 mmol/L). The most advanced model using both milk and body traits gave a slightly better fit than the model with only milk traits (R2cv = 0.63; RMSE = 0.176 mmol/L). Our findings indicate that ES of cows in early lactation can be monitored with moderately high accuracy by routine milk measurements.

Genetic background in partitioning of metabolizable energy efficiency in dairy cows.

The main objective of this study was to assess the genetic differences in metabolizable energy efficiency and efficiency in partitioning metabolizable energy in different pathways: maintenance, milk production, and growth in primiparous dairy cows. Repeatability models for residual energy intake (REI) and metabolizable energy intake (MEI) were compared and the genetic and permanent environmental variations in MEI were partitioned into its energy sinks using random regression models. We proposed 2 new feed efficiency traits: metabolizable energy efficiency (MEE), which is formed by modeling MEI fitting regressions on energy sinks [metabolic body weight (BW0.75), energy-corrected milk, body weight gain, and body weight loss] directly; and partial MEE (pMEE), where the model for MEE is extended with regressions on energy sinks nested within additive genetic and permanent environmental effects. The data used were collected from Luke's experimental farms Rehtijärvi and Minkiö between 1998 and 2014. There were altogether 12,350 weekly MEI records on 495 primiparous Nordic Red dairy cows from wk 2 to 40 of lactation. Heritability estimates for REI and MEE were moderate, 0.33 and 0.26, respectively. The estimate of the residual variance was smaller for MEE than for REI, indicating that analyzing weekly MEI observations simultaneously with energy sinks is preferable. Model validation based on Akaike's information criterion showed that pMEE models fitted the data even better and also resulted in smaller residual variance estimates. However, models that included random regression on BW0.75 converged slowly. The resulting genetic standard deviation estimate from the pMEE coefficient for milk production was 0.75 MJ of MEI/kg of energy-corrected milk. The derived partial heritabilities for energy efficiency in maintenance, milk production, and growth were 0.02, 0.06, and 0.04, respectively, indicating that some genetic variation may exist in the efficiency of using metabolizable energy for different pathways in dairy cows.

Genetic heterogeneity of feed intake, energy-corrected milk, and body weight across lactation in primiparous Holstein, Nordic Red, and Jersey cows.

In this study, we aimed to estimate and compare the genetic parameters of dry matter intake (DMI), energy-corrected milk (ECM), and body weight (BW) as 3 feed efficiency-related traits across lactation in 3 dairy cattle breeds (Holstein, Nordic Red, and Jersey). The analyses were based on weekly records of DMI, ECM, and BW per cow across lactation for 842 primiparous Holstein cows, 746 primiparous Nordic Red cows, and 378 primiparous Jersey cows. A random regression model was applied to estimate variance components and genetic parameters for DMI, ECM, and BW in each lactation week within each breed. Phenotypic means of DMI, ECM, and BW observations across lactation showed to be in very similar patterns between breeds, whereas breed differences lay in the average level of DMI, ECM, and BW. Generally, for all studied breeds, the heritability for DMI ranged from 0.2 to 0.4 across lactation and was in a range similar to the heritability for ECM. The heritability for BW ranged from 0.4 to 0.6 across lactation, higher than the heritability for DMI or ECM. Among the studied breeds, the heritability estimates for DMI shared a very similar range between breeds, whereas the heritability estimates for ECM tended to be different between breeds. For BW, the heritability estimates also tended to follow a similar range between breeds. Among the studied traits, the genetic variance and heritability for DMI varied across lactation, and the genetic correlations between DMI at different lactation stages were less than unity, indicating a genetic heterogeneity of feed intake across lactation in dairy cattle. In contrast, BW was the most genetically consistent trait across lactation, where BW among all lactation weeks was highly correlated. Genetic correlations between DMI, ECM, and BW changed across lactation, especially in early lactation. Energy-corrected milk had a low genetic correlation with both DMI and BW at the beginning of lactation, whereas ECM was highly correlated with DMI in mid and late lactation. Based on our results, genetic heterogeneity of DMI, ECM, and BW across lactation generally was observed in all studied dairy breeds, especially for DMI, which should be carefully considered for the recording strategy of these traits. The genetic correlations between DMI, ECM, and BW changed across lactation and followed similar patterns between breeds.

Detection of evaluation bias caused by genomic preselection.

The aim of this simulation study was to investigate whether it is possible to detect the effect of genomic preselection on Mendelian sampling (MS) means or variances obtained by the MS validation test. Genomic preselection of bull calves is 1 additional potential source of bias in international evaluations unless adequately accounted for in national evaluations. Selection creates no bias in traditional breeding value evaluation if the data of all animals are included. However, this is not the case with genomic preselection, as it excludes culled bulls. Genomic breeding values become biased if calculated using a multistep procedure instead of, for example, a single-step method. Currently, about 60% of the countries participating in international bull evaluations have already adopted genomic selection in their breeding schemes. The data sent for multiple across-country evaluation can, therefore, be very heterogeneous, and a proper validation method is needed to ensure a fair comparison of the bulls included in international genetic evaluations. To study the effect of genomic preselection, we generated a total of 50 replicates under control and genomic preselection schemes using the structures of the real data and pedigree from a medium-size cow population. A genetic trend of 15% of the genetic standard deviation was created for both schemes. In carrying out the analyses, we used 2 different heritabilities: 0.25 and 0.10. From the start of genomic preselection, all bulls were genomically preselected. Their MS deviations were inflated with a value corresponding to selection of the best 10% of genomically tested bull calves. For cows, the MS deviations were unaltered. The results revealed a clear underestimation of bulls' breeding values (BV) after genomic preselection started, as well as a notable deviation from zero both in true and estimated MS means. The software developed recently for the MS validation test already produces yearly MS means, and they can be used to devise an appropriate test. Mean squared true MS of genomically preselected bulls was clearly inflated. After correcting for the simulated preselection bias, the true genetic variance was smaller than the parametric value used to simulate BV, and also below the variance based on the estimated BV. Based on this study, the lower the trait's heritability, the stronger the bias in estimated BV and MS means and variances. Daughters of genomically preselected bulls had higher true and estimated BV compared with the control scheme and only slightly elevated MS means, but no effect on genetic variances was observed.

Validation of consistency of Mendelian sampling variance.

Experiences from international sire evaluation indicate that the multiple-trait across-country evaluation method is sensitive to changes in genetic variance over time. Top bulls from birth year classes with inflated genetic variance will benefit, hampering reliable ranking of bulls. However, none of the methods available today enable countries to validate their national evaluation models for heterogeneity of genetic variance. We describe a new validation method to fill this gap comprising the following steps: estimating within-year genetic variances using Mendelian sampling and its prediction error variance, fitting a weighted linear regression between the estimates and the years under study, identifying possible outliers, and defining a 95% empirical confidence interval for a possible trend in the estimates. We tested the specificity and sensitivity of the proposed validation method with simulated data using a real data structure. Moderate (M) and small (S) size populations were simulated under 3 scenarios: a control with homogeneous variance and 2 scenarios with yearly increases in phenotypic variance of 2 and 10%, respectively. Results showed that the new method was able to estimate genetic variance accurately enough to detect bias in genetic variance. Under the control scenario, the trend in genetic variance was practically zero in setting M. Testing cows with an average birth year class size of more than 43,000 in setting M showed that tolerance values are needed for both the trend and the outlier tests to detect only cases with a practical effect in larger data sets. Regardless of the magnitude (yearly increases in phenotypic variance of 2 or 10%) of the generated trend, it deviated statistically significantly from zero in all data replicates for both cows and bulls in setting M. In setting S with a mean of 27 bulls in a year class, the sampling error and thus the probability of a false-positive result clearly increased. Still, overall estimated genetic variance was close to the parametric value. Only rather strong trends in genetic variance deviated statistically significantly from zero in setting S. Results also showed that the new method was sensitive to the quality of the approximated reliabilities of breeding values used in calculating the prediction error variance. Thus, we recommend that only animals with a reliability of Mendelian sampling higher than 0.1 be included in the test and that low heritability traits be analyzed using bull data sets only.

Neglect of lactation stage leads to naive assessment of residual feed intake in dairy cattle.

Residual feed intake (RFI) is a candidate trait for feed efficiency in dairy cattle. We investigated the influence of lactation stage on the effect of energy sinks in defining RFI and the genetic parameters for RFI across lactation stages for primiparous dairy cattle. Our analysis included 747 primiparous Holstein cows, each with recordings on dry matter intake (DMI), milk yield, milk composition, and body weight (BW) over 44 lactation weeks. For each individual cow, energy-corrected milk (ECM), metabolic BW (MBW), and change in BW (ΔBW) were calculated in each week of lactation and were taken as energy sinks when defining RFI. Two RFI models were considered in the analyses; RFI model [1] was a 1-step RFI model with constant partial regression coefficients of DMI on energy sinks (ECM, MBW, and ΔBW) over lactation. In RFI model [2], data from 44 lactation weeks were divided into 11 consecutive lactation periods of 4 wk in length. The RFI model [2] was identical to model [1] except that period-specific partial regressions of DMI on ECM, MBW, and ΔBW in each lactation period were allowed across lactation. We estimated genetic parameters for RFI across lactation by both models using a random regression method. Using RFI model [2], we estimated the period-specific effects of ECM, MBW, and ΔBW on DMI in all lactation periods. Based on results from RFI model [2], the partial regression coefficients of DMI on ECM, MBW, and ΔBW differed across lactation in RFI. Constant partial regression coefficients of DMI on energy sinks over lactation was not always sufficient to account for the effects across lactation and tended to give roughly average information from all period-specific effects. Heritability for RFI over 44 lactation weeks ranged from 0.10 to 0.29 in model [1] and from 0.10 to 0.23 in model [2]. Genetic variance and heritability estimates for RFI from model [2] tended to be slightly lower and more stable across lactation than those from model [1]. In both models, RFI was genetically different over lactation, especially between early and later lactation stages. Genetic correlation estimates for RFI between early and later lactation tended to be higher when using model [2] compared with model [1]. In conclusion, partial regression coefficients of DMI on energy sinks differed across lactation when modeling RFI. Neglect of lactation stage when defining RFI could affect the assessment of RFI and the estimation of genetic parameters for RFI across lactation.

Genetic parameters for dry matter intake in primiparous Holstein, Nordic Red, and Jersey cows in the first half of lactation.

Dry matter intake (DMI) is a key component of feed efficiency in dairy cattle. In this study, we estimated genetic parameters of DMI over the first 24 lactation weeks in 3 dairy cattle breeds: Holstein, Nordic Red, and Jersey. In total, 1,656 primiparous cows (717 Holstein, 663 Nordic Red, and 276 Jersey) from Denmark, Finland, and Sweden were studied. For each breed, variance components, heritability, and repeatability for weekly DMI were estimated in 6 consecutive periods of the first 24 lactation weeks based on a repeatability animal model. Genetic correlations for DMI between different lactation periods were estimated using bivariate models. Based on our results, Holstein and Nordic Red cows had similar DMI at the beginning of lactation, but later in lactation Holstein cows had a slightly higher DMI than Nordic Red cows. In comparison, Jersey cows had a significantly lower DMI than the other 2 breeds within the first 24 lactation weeks. Heritability estimates for DMI ranged from 0.20 to 0.40 in Holsteins, 0.25 to 0.41 in Nordic Red, and 0.17 to 0.42 in Jerseys within the first 24 lactation weeks. Genetic and phenotypic variances for DMI varied along lactation within each breed and tended to be higher in the middle of lactation than at the beginning of the lactation. High genetic correlations were noted for DMI in lactation wk 5 to 24 in all 3 breeds, whereas DMI at early lactation (lactation wk 1 to 4) tended to be genetically different from DMI in the middle of lactation. The 3 breeds in this study might differ in their genetic variances for DMI, but the differences were not statistically significant in most of the studied periods. Breed differences for the genetic variance tended to be more obvious than for heritability. The potential breed differences in genetic variation for DMI should be considered in a future study using feed intake information from multiple breeds.

Cow-specific diet digestibility predictions based on near-infrared reflectance spectroscopy scans of faecal samples.

This study was designed to obtain information on prediction of diet digestibility from near-infrared reflectance spectroscopy (NIRS) scans of faecal spot samples from dairy cows at different stages of lactation and to develop a faecal sampling protocol. NIRS was used to predict diet organic matter digestibility (OMD) and indigestible neutral detergent fibre content (iNDF) from faecal samples, and dry matter digestibility (DMD) using iNDF in feed and faecal samples as an internal marker. Acid-insoluble ash (AIA) as an internal digestibility marker was used as a reference method to evaluate the reliability of NIRS predictions. Feed and composite faecal samples were collected from 44 cows at approximately 50, 150 and 250 days in milk (DIM). The estimated standard deviation for cow-specific organic matter digestibility analysed by AIA was 12.3 g/kg, which is small considering that the average was 724 g/kg. The phenotypic correlation between direct faecal OMD prediction by NIRS and OMD by AIA over the lactation was 0.51. The low repeatability and small variability estimates for direct OMD predictions by NIRS were not accurate enough to quantify small differences in OMD between cows. In contrast to OMD, the repeatability estimates for DMD by iNDF and especially for direct faecal iNDF predictions were 0.32 and 0.46, respectively, indicating that developing of NIRS predictions for cow-specific digestibility is possible. A data subset of 20 cows with daily individual faecal samples was used to develop an on-farm sampling protocol. Based on the assessment of correlations between individual sample combinations and composite samples as well as repeatability estimates for individual sample combinations, we found that collecting up to three individual samples yields a representative composite sample. Collection of samples from all the cows of a herd every third month might be a good choice, because it would yield a better accuracy.

Comparison of multiplicative heterogeneous variance adjustment models for genetic evaluations.

Two heterogeneous variance adjustment methods and two variance models were compared in a simulation study. The method used for heterogeneous variance adjustment in the Nordic test-day model, which is a multiplicative method based on Meuwissen (J. Dairy Sci., 79, 1996, 310), was compared with a restricted multiplicative method where the fixed effects were not scaled. Both methods were tested with two different variance models, one with a herd-year and the other with a herd-year-month random effect. The simulation study was built on two field data sets from Swedish Red dairy cattle herds. For both data sets, 200 herds with test-day observations over a 12-year period were sampled. For one data set, herds were sampled randomly, while for the other, each herd was required to have at least 10 first-calving cows per year. The simulations supported the applicability of both methods and models, but the multiplicative mixed model was more sensitive in the case of small strata sizes. Estimation of variance components for the variance models resulted in different parameter estimates, depending on the applied heterogeneous variance adjustment method and variance model combination. Our analyses showed that the assumption of a first-order autoregressive correlation structure between random-effect levels is reasonable when within-herd heterogeneity is modelled by year classes, but less appropriate for within-herd heterogeneity by month classes. Of the studied alternatives, the multiplicative method and a variance model with a random herd-year effect were found most suitable for the Nordic test-day model for dairy cattle evaluation.

Longitudinal modeling of residual carbon dioxide and residual feed intake in the Nordic Red dairy cattle.

Feed utilization efficiency is an important trait in dairy production playing a significant role in reducing feed costs and lowering methane emission. One of the metrics used to measure feed efficiency in dairy cows is residual feed intake (RFI). This metric requires routine measurement of feed intake. Since there is a positive high correlation between heat production and carbon dioxide (CO2) production on the one hand and heat production and efficiency on the other hand, residual carbon dioxide (RCO2) might be a useful metric to improve feed efficiency. The objectives of this study were to model the trajectories of RCO2 and RFI as well as to estimate their repeatabilities and correlations at different stages of lactation. Daily CO2 output and feed intake were recorded from 46 primiparous Nordic Red dairy cows using two Greenfeed Emissions Monitoring™ systems from 2 to 305 days in milk (DIM). Edited data comprised 5 995 daily averages. To calculate predicted values of CO2 and DM intake (DMI), prediction models were developed by fitting multiple regression models to observations. Subsequently, RCO2 and RFI were calculated by subtracting predicted values of CO2 and DMI from their corresponding actual observations. A random regression bivariate model was fitted to estimate repeatabilities and animal correlations within lactation at different DIMs between RCO2 and RFI traits. The model fitted included fixed effects of year-month of recording, lactation month, fixed regressions as well as random regressions for the animal effect. The residual variance was considered to be heterogeneous. Repeatabilities and animal correlations of RCO2 and RFI between selected DIM (for every 30 DIM i.e., 6, 36,…, 246 and 276) were calculated. Repeatability of RCO2 was high at the beginning of lactation (0.72 at DIM 6) and decreased around the peak of milk production (0.27 at DIM 96) and again increased gradually toward the end of lactation. Similarly, RFI also had high repeatability at the beginning (0.86 at DIM 6); however, it decreased in mid-lactation (0.37 at DIM 156) and then increased toward the end of lactation. Animal correlations between RCO2 and RFI were moderate to high on the same DIM and ranged from 0.37 to 0.88. Overall, we found that animals with higher CO2 production than expected also consume more DMI than expected, but the moderate correlation between RCO2 and RFI found in this study calls for more research to assess the potential of RCO2 to become a new feed efficiency metric.

Estimating breeding values for feed efficiency in dairy cattle by regression on expected feed intake.

The efficiency with which a dairy cow utilises feed for the various physiological and metabolic processes can be evaluated by metrics that contrast realised feed intake with expected feed intake. In this study, we presented a new metric - regression on expected feed intake (ReFI). This metric is based on the idea of regressing DM intake (DMI) on expected DMI using a random regression model, where energy requirement formulations are applied for the calculation of expected DMI covariables. We compared this new metric with the metrics residual feed intake (RFI) and genetic residual feed intake (gRFI), by applying them on 18 581 feed efficiency records from 654 primiparous Nordic Red dairy cows. We estimated variance components for the three metrics and their respective genetic correlations with intake and production traits. In addition, we examined the phenotypes of superior cows. With ReFI, we estimated for feed efficiency a higher genetic variation (4.7%) and heritability (0.23) compared to applying RFI or gRFI. The ReFI metric was genetically uncorrelated with DMI and negatively correlated within energy-corrected milk (ECM), whereas the RFI metric was genetically positively correlated with DMI and metabolic BW. The gRFI metric was genetically positively correlated with DMI and uncorrelated with energy sink traits. Overall, the estimated SE were large. The ReFI metric resulted in a different ranking of cows compared to those based on RFI or gRFI and was superior in selecting the most efficient animals. When the selection was based on ReFI breeding values, then the 10% most efficient cows produced 12.3% more ECM per unit metabolisable energy intake, whereas the corresponding values were only 4.3 or 5.9% when using RFI or gRFI breeding values, respectively. Based on ReFI, superior cows had also higher milk production, whereas based on RFI or gRFI milk production either decreased or was unaffected, respectively. The superiority of the ReFI metric in selecting efficient cows was due to a better modelling of the expected feed intake. The ReFI metric simplified modelling of feed utilisation efficiency in dairy cattle and resulted in breeding values that are equal to percentages of feed saved.

Employing a Monte Carlo algorithm in expectation maximization restricted maximum likelihood estimation of the linear mixed model.

Multiple-trait and random regression models have multiplied the number of equations needed for the estimation of variance components. To avoid inversion or decomposition of a large coefficient matrix, we propose estimation of variance components by Monte Carlo expectation maximization restricted maximum likelihood (MC EM REML) for multiple-trait linear mixed models. Implementation is based on full-model sampling for calculating the prediction error variances required for EM REML. Performance of the analytical and the MC EM REML algorithm was compared using a simulated and a field data set. For field data, results from both algorithms corresponded well even with one MC sample within an MC EM REML round. The magnitude of the standard errors of estimated prediction error variances depended on the formula used to calculate them and on the MC sample size within an MC EM REML round. Sampling variation in MC EM REML did not impair the convergence behaviour of the solutions compared with analytical EM REML analysis. A convergence criterion that takes into account the sampling variation was developed to monitor convergence for the MC EM REML algorithm. For the field data set, MC EM REML proved far superior to analytical EM REML both in computing time and in memory need.

Animal board invited review: Genomic-based improvement of cattle in response to climate change.

Climate change brings challenges to cattle production, such as the need to adapt to new climates and pressure to reduce greenhouse emissions (GHG). In general, the improvement of traits in current breeding goals is favourably correlated with the reduction of GHG. Current breeding goals and tools for increasing cattle production efficiency have reduced GHG. The same amount of production can be achieved by a much smaller number of animals. Genomic selection (GS) may offer a cost-effective way of using an efficient breeding approach, even in low- and middle-income countries. As climate change increases the intensity of heatwaves, adaptation to heat stress leads to lower efficiency of production and, thus, is unfavourable to the goal of reducing GHG. Furthermore, there is evidence that heat stress during cow pregnancy can have many generation-long lowering effects on milk production. Both adaptation and reduction of GHG are among the difficult-to-measure traits for which GS is more efficient and suitable than the traditional non-genomic breeding evaluation approach. Nevertheless, the commonly used within-breed selection may be insufficient to meet the new challenges; thus, cross-breeding based on selecting highly efficient and highly adaptive breeds may be needed. Genomic introgression offers an efficient approach for cross-breeding that is expected to provide high genetic progress with a low rate of inbreeding. However, well-adapted breeds may have a small number of animals, which is a source of concern from a genetic biodiversity point of view. Furthermore, low animal numbers also limit the efficiency of genomic introgression. Sustainable cattle production in countries that have already intensified production is likely to emphasise better health, reproduction, feed efficiency, heat stress and other adaptation traits instead of higher production. This may require the application of innovative technologies for phenotyping and further use of new big data techniques to extract information for breeding.

Marker-assisted breeding value estimation for mastitis resistance in Finnish Ayrshire cattle.

Marker-assisted breeding value estimation is expected to increase the accuracy of estimated breeding values, especially for traits with low heritability. Several quantitative trait loci (QTL) have been found for somatic cell score and clinical mastitis. The objective of this study was to demonstrate marker-assisted breeding value estimation, combining data of genotyped and ungenotyped animals in a large pedigree population using either identical-by-descent (IBD) or identical-by-state (IBS) haplotypes for some previously identified QTL regions for somatic cell score and clinical mastitis in Finnish Ayrshire cattle. For both methods, QTL variances were estimated based on daughter yield deviations of genotyped bulls. The QTL explained only a small proportion of genetic variance, especially with IBS haplotypes. Using IBD haplotypes gave more reranking of bulls and cows than using IBS haplotypes. Cross-validation showed no increase in predictive ability when using IBS haplotypes compared with conventional breeding value estimation, whereas a decrease in predictive ability was observed with IBD haplotypes. Furthermore, computing time was lower and convergence was better with IBS haplotypes than with IBD haplotypes. In this study on mastitis resistance in Finnish Ayrshire, conventional breeding value estimation would be advocated because of the lack in improvement of accuracy and predictive ability when using marker-assisted breeding value estimation. However, in situations where IBS haplotypes would explain 10 to 20% or more of the genetic variance, marker-assisted breeding value estimation with IBS haplotypes may yield greater accuracy and predictive ability than conventional breeding value estimation.

Estimation of social genetic effects on feeding behaviour and production traits in pigs.

Pigs are housed in groups during the test period. Social effects between pen mates may affect average daily gain (ADG), backfat thickness (BF), feed conversion rate (FCR), and the feeding behaviour traits of pigs sharing the same pen. The aim of our study was to estimate the genetic parameters of feeding behaviour and production traits with statistical models that include social genetic effects (SGEs). The data contained 3075 Finnish Yorkshire, 3351 Finnish Landrace, and 968 F1-crossbred pigs. Feeding behaviour traits were measured as the number of visits per day (NVD), time spent in feeding per day (TPD), daily feed intake (DFI), time spent in feeding per visit (TPV), feed intake per visit (FPV), and feed intake rate (FR). The test period was divided into five periods of 20 days. The number of pigs per pen varied from 8 to 12. Two model approaches were tested, i.e. a fixed group size model and a variable group size model. For the fixed group size model, eight random pigs per pen were included in the analysis, while all pigs in a pen were included for the variable group size model. The linear mixed-effects model included sex, breed, and herd*year*season as fixed effects and group (batch*pen), litter, the animal itself (direct genetic effect (DGE)), and pen mates (SGEs) as random effects. For feeding behaviour traits, estimates of the total heritable variation (T2± SE) and classical heritability (h2± SE, values given in brackets) from the variable group size model (e.g. period 1) were 0.34 ± 0.13 (0.30 ± 0.04) for NVD, 0.41 ± 0.10 (0.37 ± 0.04) for TPD, 0.40 ± 0.15 (0.14 ± 0.03) for DFI, 0.53 ± 0.15 (0.28 ± 0.04) for TPV, 0.66 ± 0.17 (0.28 ± 0.04) for FPV, and 0.29 ± 0.13 (0.22 ± 0.03) for FR. The effect of social interaction was minimal for ADG (T2 = 0.29 ± 0.11 and h2 = 0.29 ± 0.04), BF (T2 = 0.48 ± 0.12 and h2 = 0.38 ± 0.07), and FCR (T2 = 0.37 ± 0.12 and h2 = 0.29 ± 0.04) using the variable group size model. In conclusion, the results indicate that social interactions have a considerable indirect genetic effect on the feeding behaviour and FCR of pigs but not on ADG and BF.

Genetic analyses of metabolic body weight, carcass weight and body conformation traits in Nordic dairy cattle.

Improving feed efficiency in dairy cattle by animal breeding has started in the Nordic countries. One of the two traits included in the applied Saved feed index is called maintenance and it is based on the breeding values for metabolic BW (MBW). However, BW recording based on heart girth measurements is decreasing and recording based on scales is increasing only slowly, which may weaken the maintenance index in future. Therefore, the benefit of including correlated traits, like carcass weight and conformation traits, is of interest. In this study, we estimated genetic variation and genetic correlations for eight traits describing the energy requirement for maintenance in dairy cattle including: first, second and third parity MBW based on heart girth measurements, carcass weight (CARW) and predicted MBW (pMBW) based on predicted slaughter weight, and first parity conformation traits stature (ST), chest width (CW) and body depth (BD). The data consisted of 21329 records from Finnish Ayrshire and 9780 records from Holstein cows. Heritability estimates were 0.44, 0.53, 0.56, 0.52, 0.54, 0.60, 0.17 and 0.26 for MBW1, MBW2, MBW3, CARW, pMBW, ST, CW and BD, respectively. Estimated genetic correlations among MBW traits were strong (>0.95). Genetic correlations between slaughter traits (CARW and pMBW) and MBW traits were higher (from 0.77 to 0.90) than between conformation and MBW traits (from 0.47 to 0.70). Our results suggest that including information on carcass weight and body conformation as correlated traits into the maintenance index is beneficial when direct BW measurements are not available or are difficult or expensive to obtain.

Genetic correlations between energy status indicator traits and female fertility in primiparous Nordic Red Dairy cattle.

Inclusion of feed efficiency traits into the dairy cattle breeding programmes will require considering early lactation energy status to avoid deterioration in health and fertility of dairy cows. In this regard, energy status indicator (ESI) traits, for example, blood metabolites or milk fatty acids (FAs), are of interest. These indicators can be predicted from routine milk samples by mid-IR reflectance spectroscopy (MIR). In this study, we estimated genetic variation in ESI traits and their genetic correlation with female fertility in early lactation. The data consisted of 37 424 primiparous Nordic Red Dairy cows with milk test-day records between 8 and 91 days in milk (DIM). Routine test-day milk samples were analysed by MIR using previously developed calibration equations for blood plasma non-esterified FA (NEFA), milk FAs, milk beta-hydroxybutyrate (BHB) and milk acetone concentrations. Six ESI traits were considered and included: plasma NEFA concentration (mmol/l) either predicted by multiple linear regression including DIM, milk fat to protein ratio (FPR) and FAs C10:0, C14:0, C18:1 cis-9, C14:0 * C18:1 cis-9 (NEFAFA) or directly from milk MIR spectra (NEFAMIR), C18:1 cis-9 (g/100 ml milk), FPR, BHB (mmol/l milk) and acetone (mmol/l milk). The interval from calving to first insemination (ICF) was considered as the fertility trait. Data were analysed using linear mixed models. Heritability estimates varied during the first three lactation months from 0.13 to 0.19, 0.10 to 0.17, 0.09 to 0.14, 0.07 to 0.10, 0.13 to 0.17 and 0.13 to 0.18 for NEFAMIR, NEFAFA, C18:1 cis-9, FPR, milk BHB and acetone, respectively. Genetic correlations between all ESI traits and ICF were from 0.18 to 0.40 in the first lactation period (8 to 35 DIM), in general somewhat lower (0.03 to 0.43) in the second period (36 to 63 DIM) and decreased clearly (-0.02 to 0.19) in the third period (64 to 91 DIM). Our results indicate that genetic variation in energy status of cows in early lactation can be determined using MIR-predicted indicators. In addition, the markedly lower genetic correlation between ESI traits and fertility in the third lactation month indicated that energy status should be determined from the first test-day milk samples during the first 2 months of lactation.

Review: Selecting for improved feed efficiency and reduced methane emissions in dairy cattle.

It may be possible for dairy farms to improve profitability and reduce environmental impacts by selecting for higher feed efficiency and lower methane (CH4) emission traits. It remains to be clarified how CH4 emission and feed efficiency traits are related to each other, which will require direct and accurate measurements of both of these traits in large numbers of animals under the conditions in which they are expected to perform. The ranking of animals for feed efficiency and CH4 emission traits can differ depending upon the type and duration of measurement used, the trait definitions and calculations used, the period in lactation examined and the production system, as well as interactions among these factors. Because the correlation values obtained between feed efficiency and CH4 emission data are likely to be biased when either or both are expressed as ratios, therefore researchers would be well advised to maintain weighted components of the ratios in the selection index. Nutrition studies indicate that selecting low emitting animals may result in reduced efficiency of cell wall digestion, that is NDF, a key ruminant characteristic in human food production. Moreover, many interacting biological factors that are not measured directly, including digestion rate, passage rate, the rumen microbiome and rumen fermentation, may influence feed efficiency and CH4 emission. Elucidating these mechanisms may improve dairy farmers ability to select for feed efficiency and reduced CH4 emission.