Short communication: Genetic variation of saturated fatty acids in Holsteins in the Walloon region of Belgium

Random regression test-day models using Legendre polynomials are commonly used for the estimation of genetic parameters and genetic evaluation for test-day milk production traits. However, some researchers have reported that these models present some undesirable properties such as the overestimation of variances at the edges of lactation. Describing genetic variation of saturated fatty acids expressed in milk fat might require the testing of different models. Therefore, 3 different functions were used and compared to take into account the lactation curve: (1) Legendre polynomials with the same order as currently applied for genetic model for production traits; 2) linear splines with 10 knots; and 3) linear splines with the same 10 knots reduced to 3 parameters. The criteria used were Akaike's information and Bayesian information criteria, percentage square biases, and log-likelihood function. These criteria indentified Legendre polynomials and linear splines with 10 knots reduced to 3 parameters models as the most useful. Reducing more complex models using eigenvalues seemed appealing because the resulting models are less time demanding and can reduce convergence difficulties, because convergence properties also seemed to be improved. Finally, the results showed that the reduced spline model was very similar to the Legendre polynomials model.     

Use of test-day records beyond three hundred five days for estimation of three hundred five-day breeding values for production traits and somatic cell score of Canadian Holsteins

The Canadian Test-Day Model includes test-day (TD) records from 5 to 305 d in milk (DIM). Because 60% of Canadian Holstein cows have at least one lactation longer than 305 d, a significant number of TD records beyond 305 DIM could be included in the genetic evaluation. The aim of this study was to investigate whether TD records beyond 305 DIM could be useful for estimation of 305-d estimated breeding value (EBV) for milk, fat, and protein yields and somatic cell score. Data were 48,638,184 TD milk, fat, and protein yields and somatic cell scores from the first 3 lactations of 2,826,456 Canadian Holstein cows. All production traits were preadjusted for the effect of pregnancy. Subsets of data were created for variance-component estimation by random sampling of 50 herds. Variance components were estimated using Gibbs sampling. Full data sets were used for estimation of breeding values. Three multiple-trait, multiple-lactation random regression models with TD records up to 305 DIM (M305), 335 DIM (M335), and 365 DIM (M365) were fitted. Two additional models (M305a and M305b) used TD records up to 305 DIM and variance components previously estimated by M335 and M365, respectively. The effects common to all models were fixed effects of herd × test-date and DIM class, fixed regression on DIM nested within region × age × season class, and random regressions for additive genetic and permanent environmental effects. Legendre polynomials of order 6 and 4 were fitted for fixed and random regressions, respectively. Rapid increase of additive genetic and permanent environmental variances at extremes of lactations was observed with all 3 models. The increase of additive genetic and permanent environmental variances was at earlier DIM with M305, resulting in greater variances at 305 DIM with M305 than with M335 and M365. Model M305 had the best ability to predict TD yields from 5 through 305 DIM and less error of prediction of 305-d EBV than M335 and M365. Model M335 had smaller change of 305-d EBV of bulls over the period of 7 yr than did M305 and M365. Model M305a had the least error of prediction and change of 305-d EBV from all models. Therefore, the use of TD records of Holstein cows from 5 through 305 DIM and variance components estimated using records up to 335 DIM is recommended for the Canadian Test-Day Model.     

Relationships between milk yield and somatic cell score in Canadian Holsteins from simultaneous and recursive random regression models

Multiple-trait random regression animal models with simultaneous and recursive links between phenotypes for milk yield and somatic cell score (SCS) on the same test day were fitted to Canadian Holstein data. All models included fixed herd test-day effects and fixed regressions within region-age at calving-season of calving classes, and animal additive genetic and permanent environmental regressions with random coefficients. Regressions were Legendre polynomials of order 4 on a scale from 5 to 305 d in milk (DIM). Bayesian methods via Gibbs sampling were used for the estimation of model parameters. Heterogeneity of structural coefficients was modeled across (the first 3 lactations) and within (4 DIM intervals) lactation. Model comparisons in terms of Bayes factors indicated the superiority of simultaneous models over the standard multiple-trait model and recursive parameterizations. A moderate heterogeneous (both across- and within-lactation) negative effect of SCS on milk yield (from −0.36 for 116 to 265 DIM in lactation 1 to −0.81 for 5 to 45 DIM in lactation 3) and a smaller positive reciprocal effect of SCS on milk yield (from 0.007 for 5 to 45 DIM in lactation 2 to 0.023 for 46 to 115 DIM in lactation 3) were estimated in the most plausible specification. No noticeable differences among models were detected for genetic and environmental variances and genetic parameters for the first 2 regression coefficients. The curves of genetic and permanent environmental variances, heritabilities, and genetic and phenotypic correlations between milk yield and SCS on a daily basis were different for different models. Rankings of bulls and cows for 305-d milk yield, average daily SCS, and milk lactation persistency remained the same among models. No apparent benefits are expected from fitting causal phenotypic relationships between milk yield and SCS on the same test day in the random regression test-day model for genetic evaluation purposes.     

Joint international evaluation of Milking Shorthorn dairy cattle for production traits

Pedigree information and test-day records for the first 3 parities of Milking Shorthorn dairy cattle from 5 countries were analyzed. After editing, the data included 1,018,528 test-day records from 68,653 cows. A multiple-lactation random regression test-day model with Legendre polynomials of order 4 and a Bayesian method were used to estimate variance components for both single and multiple-countries. Fixed effects included herd-test-day class and regressions on DIM within age at calving-parity-season of calving. Random effects included animal genetic, permanent environmental, and residual effects. Average daily heritabilities from single country analyses ranged from 0.33 to 0.47 for milk yield and from 0.37 to 0.45 for protein yield across lactations and countries. Common sires (66) and their daughters were identified for creating a connected data set for simultaneous (co)variance component estimation of milk yield across all 5 countries. Between-country genetic correlations were low, with values from 0.08 to 0.46 and standard deviations from 0.08 to 0.12. Estimated breeding values for milk were generated for each animal using the same test-day animal model. Correlations among country estimated breeding values were higher than genetic correlations. Top 100 bull lists were generated on the scale of each country, and genetic progress was assessed. Future evaluation with increased genetic ties among countries may facilitate international comparison of Milking Shorthorns.     

Random herd curves in a test-day model for milk, fat, and protein production of dairy cattle in The Netherlands

With random regression models, genetic parameters of test-day milk production records of dairy cattle can be estimated directly from the data. However, several researchers that used this method have reported unrealistically high variances at the borders of the lactation trajectory and low genetic correlations between beginning and end of lactation. Recently, it has been proposed to include herd-specific regression curves in the random regression model. The objective was to study the effect of including random herd curves on estimated genetic parameters. Genetic parameters were estimated with 2 models; both included random regressions for the additive genetic and permanent environmental effect, whereas the second model also included a random regression effect for herd x 2-yr period of calving. All random regressions were modeled with fourth-order Legendre polynomials. Bayesian techniques with Gibbs sampling were used to estimate all parameters. The data set comprised 857,255 test-day milk, fat, and protein records from lactations 1, 2, and 3 of 43,990 Holstein cows from 544 herds. Genetic variances estimated by the second model were lower in the first 100 d and at the end of the lactation, especially in lactations 2 and 3. Genetic correlations between d 50 and the end of lactation were around 0.25 higher in the second model and were consistent with studies where lactation stages are modeled as different traits. Subsequently, estimated heritabilities for persistency were up to 0.14 lower in the second model. It is suggested to include herd curves in a random regression model when estimating genetic parameters of test-day production traits in dairy cattle.     

Detection of different shapes of lactation curve for milk yield in dairy cattle by empirical mathematical models

The study of relationships between mathematical properties of functions used to model lactation curves is usually limited to the evaluation of the goodness of fit. Problems related to the existence of different lactation curve shapes are usually neglected or solved drastically by considering shapes markedly different from the standard as biologically atypical. A deeper investigation could yield useful indications for developing technical tools aimed at modifying the lactation curve in a desirable fashion. Relationships between mathematical properties and lactation curve shapes were analyzed by fitting several common functions (Wood incomplete gamma, Wilmink's exponential, Ali and Schaeffer's polynomial regression, and fifth-order Legendre polynomials) to 229,518 test-day records belonging to 27,837 lactations of Italian Simmental cows. Among the best fits (adjusted r(2) higher than 0.75), the 3-parameter models (Wood and Wilmink) were able to detect 2 main groups of curve shape: standard and atypical. Five-parameter models (Ali and Schaeffer function and the Legendre polynomials) were able to recognize a larger number of curve shapes. The higher flexibility of 5-parameter models was accompanied by increased sensitivity to local random variation as evidenced by the bias in estimated test-day yields at the beginning and end of lactation (border effect). Meaning of parameters, range of their values and of their (co) variances are clearly different among groups of curves. Our results suggest that analysis based on comparisons between parameter values and (co)variances should be done carefully. Comparisons among parameter values and (co)variances could yield more robust, reliable, and easy to interpret results if performed within groups based on curve shape.     

Parametric correlation functions to model the structure of permanent environmental (co)variances in milk yield random regression models

The objective of the present study was to estimate milk yield genetic parameters applying random regression models and parametric correlation functions combined with a variance function to model animal permanent environmental effects. A total of 152,145 test-day milk yields from 7,317 first lactations of Holstein cows belonging to herds located in the southeastern region of Brazil were analyzed. Test-day milk yields were divided into 44 weekly classes of days in milk. Contemporary groups were defined by herd-test-day comprising a total of 2,539 classes. The model included direct additive genetic, permanent environmental, and residual random effects. The following fixed effects were considered: contemporary group, age of cow at calving (linear and quadratic regressions), and the population average lactation curve modeled by fourth-order orthogonal Legendre polynomial. Additive genetic effects were modeled by random regression on 1/6/2010orthogonal Legendre polynomials of days in milk, whereas permanent environmental effects were estimated using a stationary or nonstationary parametric correlation function combined with a variance function of different orders. The structure of residual variances was modeled using a step function containing 6 variance classes. The genetic parameter estimates obtained with the model using a stationary correlation function associated with a variance function to model permanent environmental effects were similar to those obtained with models employing orthogonal Legendre polynomials for the same effect. A model using a sixth-order polynomial for additive effects and a stationary parametric correlation function associated with a seventh-order variance function to model permanent environmental effects would be sufficient for data fitting.     

Genetic evaluation of bulls and cows with single- and multiple-country test-day models

First-lactation milk yield test-day records on cows from Australia, Canada, Italy, and New Zealand were analyzed by single- and multiple-country random regression models. Models included fixed effects of herd-test day and breed composition-age at calving-season of calving by days in milk, and random regressions with Legendre polynomials of order four for animal genetic and permanent environmental effects. Milk yields in different countries were defined as genetically different traits for the purpose of multiple-trait model. Estimated breeding values of bulls and cows from single- and multiple-trait models were compared within and across countries for two traits: total milk yield in lactation and lactation persistency, defined as the linear coefficient of animal genetic curve. Correlations between single- and multiple-trait evaluations within country for total yield were higher than 0.95 for bulls and close to 1 for cows. Correlations for lactation persistency were lower than respective correlations for total yield. Between country correlations for lactation yield ranged from 0.93 to 0.96, indicating different ranking of bulls on different country scales under multiple-trait model. Lactation persistency had in general lower between-country correlations, with the highest values for Canada-Italy and Australia-New Zealand pairs, for both single- and multiple-country models. Although multiple-country random regression test-day model was computationally feasible for four countries, the same would not be true for routine international genetic evaluation in the near future.     

Genetic analyses of protein yield in dairy cows applying random regression models with time-dependent and temperature x humidity-dependent covariates

Data used in the present study included 1,095,980 first-lactation test-day records for protein yield of 154,880 Holstein cows housed on 196 large-scale dairy farms in Germany. Data were recorded between 2002 and 2009 and merged with meteorological data from public weather stations. The maximum distance between each farm and its corresponding weather station was 50 km. Hourly temperature-humidity indexes (THI) were calculated using the mean of hourly measurements of dry bulb temperature and relative humidity. On the phenotypic scale, an increase in THI was generally associated with a decrease in daily protein yield. For genetic analyses, a random regression model was applied using time-dependent (d in milk, DIM) and THI-dependent covariates. Additive genetic and permanent environmental effects were fitted with this random regression model and Legendre polynomials of order 3 for DIM and THI. In addition, the fixed curve was modeled with Legendre polynomials of order 3. Heterogeneous residuals were fitted by dividing DIM into 5 classes, and by dividing THI into 4 classes, resulting in 20 different classes. Additive genetic variances for daily protein yield decreased with increasing degrees of heat stress and were lowest at the beginning of lactation and at extreme THI. Due to higher additive genetic variances, slightly higher permanent environment variances, and similar residual variances, heritabilities were highest for low THI in combination with DIM at the end of lactation. Genetic correlations among individual values for THI were generally >0.90. These trends from the complex random regression model were verified by applying relatively simple bivariate animal models for protein yield measured in 2 THI environments; that is, defining a THI value of 60 as a threshold. These high correlations indicate the absence of any substantial genotype × environment interaction for protein yield. However, heritabilities and additive genetic variances from the random regression model tended to be slightly higher in the THI range corresponding to cows’ comfort zone. Selecting such superior environments for progeny testing can contribute to an accurate genetic differentiation among selection candidates.     

A general method to validate breeding value prediction software

The validity of national genetic evaluations depends on the quality of input data, on the model of analysis, and on the correctness of genetic evaluation software. A general strategy was developed to validate national breeding value prediction software: performances from a real data file were replaced with simulated ones, created from simulated fixed and random effects and residuals in such a way that BLUP estimates from the evaluation software must be equal to the simulated effects. This approach was implemented for a multiple-trait model and a random regression test-day model. An example was presented on test-day observations analyzed with a random regression animal model including a lactation curve described as a sum of fixed polynomial regression and fixed spline regression on days in milk, and with genetic and permanent environmental effects modeled by using Legendre polynomials of order 2. Residuals had heterogeneous variances, and phantom parent groups were included. This method can be easily extended to other linear models. The comparison of genetic evaluation results with simulated true effects is used to demonstrate the great efficiency and usefulness of the proposed method.     

Genetic parameter estimation for milk yield over multiple parities and various lengths of lactation in Danish Jerseys by random regression models

The objectives of this study were to test for heterogeneity of genetic and environmental variance among completed and extended records from different lactations or different days in milk (DIM) and to build a model that accounts for this heterogeneity. A total of 147,457 305-d milk yield records from Danish Jersey cows calving between 1984 and early 1999 from two regions of Denmark were used in this study. Results showed that DIM and parity influenced parameters estimated from an animal model with repeated records. Therefore, the data were analyzed using random-regression models that allow the covariance between measurements to change gradually with DIM and parity. Random regressions were fitted for additive genetic effects and permanent environmental effects using second- or third-order normalized Legendre polynomials for DIM and parity. Variances of random-regression coefficients associated with all orders of the polynomials were significant. Based on these parameter estimates, a covariance function (CF) was defined. The CF showed that the heritability decreases over parities, but within each parity heritability increases with DIM, whereas variance of permanent environmental effects increases over parities and decreases with DIM. Generally, genetic correlations were higher between records with similar DIM and parity. The results indicate that there are problems with the extension procedure used to predict 305-d milk yields. Using the covariance functions estimated in this study, breeding values could be predicted that take into account the covariance structure between records from different parities and different DIM.     

Genetic parameters of milking frequency and milk production traits in Canadian Holsteins milked by an automated milking system

Twice-a-day milking is currently the most frequently used milking schedule in Canadian dairy cattle. However, with an automated milking system (AMS), dairy cows can be milked more frequently. The objective of this study was to estimate genetic parameters for milking frequency and for production traits of cows milked within an AMS. Data were 141,927 daily records of 953 primiparous Holstein cows from 14 farms in Ontario and Quebec. Most cows visited the AMS 2 (46%) or 3 (37%) times a day. A 2-trait [daily (24-h) milking frequency and daily (24-h) milk yield] random regression daily animal model and a multiple-trait (milk, fat, protein yields, somatic cell score, and milking frequency) random regression test-day animal model were used for the estimation of (co)variance components. Both models included fixed effect of herd × test-date, fixed regressions on days in milk (DIM) nested within age at calving by season of calving, and random regressions for additive genetic and permanent environmental effects. Both fixed and random regressions were fitted with fourth-order Legendre polynomials on DIM. The number of cows in the multiple-trait test-day model was smaller compared with the daily animal model. Heritabilities from the daily model for daily (24-h) milking frequency and daily (24-h) milk yield ranged between 0.02 and 0.08 and 0.14 and 0.20, respectively. Genetic correlations between daily (24-h) milk yield and daily (24-h) milking frequency were largest at the end of lactation (0.80) and smallest in mid-lactation (0.27). Heritabilities from the test-day model for test-day milking frequency, milk, fat and protein yield, and somatic cell score were 0.14, 0.26, 0.20, 0.21, and 0.20, respectively. The genetic correlation was positive between test-day milking frequency and official test-day milk, fat, and protein yields, and negative between official test-day somatic cell score and test-day milking frequency.     

Estimation of genetic parameters for milk yield in Murrah buffaloes by Bayesian inference

Random regression models were used to estimate genetic parameters for test-day milk yield in Murrah buffaloes using Bayesian inference. Data comprised 17,935 test-day milk records from 1,433 buffaloes. Twelve models were tested using different combinations of third-, fourth-, fifth-, sixth-, and seventh-order orthogonal polynomials of weeks of lactation for additive genetic and permanent environmental effects. All models included the fixed effects of contemporary group, number of daily milkings and age of cow at calving as covariate (linear and quadratic effect). In addition, residual variances were considered to be heterogeneous with 6 classes of variance. Models were selected based on the residual mean square error, weighted average of residual variance estimates, and estimates of variance components, heritabilities, correlations, eigenvalues, and eigenfunctions. Results indicated that changes in the order of fit for additive genetic and permanent environmental random effects influenced the estimation of genetic parameters. Heritability estimates ranged from 0.19 to 0.31. Genetic correlation estimates were close to unity between adjacent test-day records, but decreased gradually as the interval between test-days increased. Results from mean squared error and weighted averages of residual variance estimates suggested that a model considering sixth- and seventh-order Legendre polynomials for additive and permanent environmental effects, respectively, and 6 classes for residual variances, provided the best fit. Nevertheless, this model presented the largest degree of complexity. A more parsimonious model, with fourth- and sixth-order polynomials, respectively, for these same effects, yielded very similar genetic parameter estimates. Therefore, this last model is recommended for routine applications.     

Modeling daily energy balance of dairy cows in the first three lactations

Daily energy balance was calculated for 111 Holstein cows in their first 3 lactations, based on combinations of smoothed preadjusted phenotypic records for milk yield, feed intake, live weight, and body condition score. Two energy balance traits were defined: one based on milk yield and feed intake (EB1) and the other on live weight and body condition score change (EB2). Bessel functions (BF), Legendre polynomials (LP), sinusoidal functions (SF), and cubic splines (CS) were used to model energy balance within and across lactations. Models with BF or LP fitted fixed regressions of order 1 to 6 and random regressions of order 1 to 10. Cubic splines were fitted at 5 to 30 equally spaced knot points. In within-lactation analyses with BF and LP models, likelihood ratio tests revealed that the fit improved significantly up to random regression order of 5 for EB1 and 4 for EB2, independently of the fixed regression order. For EB1 analyses with LP, improvement was marginal albeit significant even for higher random regression order. For CS models, optimal number of knot points was 13 and 12 for EB1 and EB2, respectively. Residual variance and comparisons between actual and predicted energy balance showed that LP of minimum order 8 and 5 modeled, respectively, EB1 and EB2 better than the other 3 functions. In across-lactation analyses with BF and LP models, likelihood ratio tests were significant as the random regression order increased, for any order of the fixed regression. For CS models, optimal number of knot points was 14 and 16 for EB1 and EB2, respectively. Residual variance and comparisons between actual and predicted energy balance showed that models fitting CS and high (>8) random order BF or LP provided the best fit to both traits. However, in an across-lactation analysis, even higher order of LP or BF will be required to provide as good a fit as within-lactation analyses.     

Multiple-trait random regression models for the estimation of genetic parameters for milk,fat, and protein yield in buffaloes

In this study, genetic parameters for test-day milk, fat, and protein yield were estimated for the first lactation. The data analyzed consisted of 1,433 first lactations of Murrah buffaloes, daughters of 113 sires from 12 herds in the state of São Paulo, Brazil, with calvings from 1985 to 2007. Ten-month classes of lactation days were considered for the test-day yields. The (co)variance components for the 3 traits were estimated using the regression analyses by Bayesian inference applying an animal model by Gibbs sampling. The contemporary groups were defined as herd-year-month of the test day. In the model, the random effects were additive genetic, permanent environment, and residual. The fixed effects were contemporary group and number of milkings (1 or 2), the linear and quadratic effects of the covariable age of the buffalo at calving, as well as the mean lactation curve of the population, which was modeled by orthogonal Legendre polynomials of fourth order. The random effects for the traits studied were modeled by Legendre polynomials of third and fourth order for additive genetic and permanent environment, respectively, the residual variances were modeled considering 4 residual classes. The heritability estimates for the traits were moderate (from 0.21–0.38), with higher estimates in the intermediate lactation phase. The genetic correlation estimates within and among the traits varied from 0.05 to 0.99. The results indicate that the selection for any trait test day will result in an indirect genetic gain for milk, fat, and protein yield in all periods of the lactation curve. The accuracy associated with estimated breeding values obtained using multi-trait random regression was slightly higher (around 8%) compared with single-trait random regression. This difference may be because to the greater amount of information available per animal.     

Comparison of random regression test-day models for Polish Black and White cattle

Test-day milk yields of first-lactation Black and White cows were used to select the model for routine genetic evaluation of dairy cattle in Poland. The population of Polish Black and White cows is characterized by small herd size, low level of production, and relatively early peak of lactation. Several random regression models for first-lactation milk yield were initially compared using the “percentage of squared bias” criterion and the correlations between true and predicted breeding values. Models with random herd-test-date effects, fixed age-season and herd-year curves, and random additive genetic and permanent environmental curves (Legendre polynomials of different orders were used for all regressions) were chosen for further studies. Additional comparisons included analyses of the residuals and shapes of variance curves in days in milk. The low production level and early peak of lactation of the breed required the use of Legendre polynomials of order 5 to describe age-season lactation curves. For the other curves, Legendre polynomials of order 3 satisfactorily described daily milk yield variation. Fitting third-order polynomials for the permanent environmental effect made it possible to adequately account for heterogeneous residual variance at different stages of lactation.     

Modeling heat stress effect on Holstein cows under hot and dry conditions: Selection tools

Data from milk recording of Holstein-Friesian cows together with weather information from 2 regions in Southern Spain were used to define the models that can better describe heat stress response for production traits and somatic cell score (SCS). Two sets of analyses were performed, one aimed at defining the population phenotypic response and the other at studying the genetic components. The first involved 2,514,762 test-day records from up to 5 lactations of 128,112 cows. Two models, one fitting a comfort threshold for temperature and a slope of decay after the threshold, and the other a cubic Legendre polynomial (LP) model were tested. Average (T_AVE) and maximum daily temperatures were alternatively considered as covariates. The LP model using T_AVE as covariate showed the best goodness of fit for all traits. Estimated rates of decay from this model for production at 25 and 34°C were 36 and 170, 3.8 and 3.0, and 3.9 and 8.2 g/d per degree Celsius for milk, fat, and protein yield, respectively. In the second set of analyses, a sample of 280,958 test-day records from first lactations of 29,114 cows was used. Random regression models including quadratic or cubic LP regressions (TEM_) on T_AVE or a fixed threshold and an unknown slope (DUMMY), including or not cubic regressions on days in milk (DIM3_), were tested. For milk and SCS, the best models were the DIM3_ models. In contrast, for fat and protein yield, the best model was TEM3. The DIM3DUMMY models showed similar performance to DIM3TEM3. The estimated genetic correlations between the same trait under cold and hot temperatures (ρ) indicated the existence of a large genotype by environment interaction for fat (ρ = 0.53 for model TEM3) and protein yield (ρ around 0.6 for DIM3TEM3) and for SCS (ρ = 0.64 for model DIM3TEM3), and a small genotype by environment interaction for milk (ρ over 0.8). The eigendecomposition of the additive genetic covariance matrix from model TEM3 showed the existence of a dominant component, a constant term that is not affected by temperature, representing from 64% of the variation for SCS to 91% of the variation for milk. The second component, showing a flat pattern at intermediate temperatures and increasing or decreasing slopes for the extremes, gathered 15, 11, and 24% of the variation for fat and protein yield and SCS, respectively. This component could be further evaluated as a selection criterion for heat tolerance independently of the production level.     

Genetic analysis of somatic cell score in Norwegian cattle using random regression test-day models

The dataset used in this analysis contained a total of 341,736 test-day observations of somatic cell scores from 77,110 primiparous daughters of 1965 Norwegian Cattle sires. Initial analyses, using simple random regression models without genetic effects, indicated that use of homogeneous residual variance was appropriate. Further analyses were carried out by use of a repeatability model and 12 random regression sire models. Legendre polynomials of varying order were used to model both permanent environmental and sire effects, as did the Wilmink function, the Lidauer-M?ntysaari function, and the Ali-Schaeffer function. For all these models, heritability estimates were lowest at the beginning (0.05 to 0.07) and higher at the end (0.09 to 0.12) of lactation. Genetic correlations between somatic cell scores early and late in lactation were moderate to high (0.38 to 0.71), whereas genetic correlations for adjacent DIM were near unity. Models were compared based on likelihood ratio tests, Bayesian information criterion, Akaike information criterion, residual variance, and predictive ability. Based on prediction of randomly excluded observations, models with 4 coefficients for permanent environmental effect were preferred over simpler models. More highly parameterized models did not substantially increase predictive ability. Evaluation of the different model selection criteria indicated that a reduced order of fit for sire effects was desireable. Models with zeroth- or first-order of fit for sire effects and higher order of fit for permanent environmental effects probably underestimated sire variance. The chosen model had Legendre polynomials with 3 coefficients for sire, and 4 coefficients for permanent environmental effects. For this model, trajectories of sire variance and heritability were similar assuming either homogeneous or heterogeneous residual variance structure.     

Estimates of genetic parameters and eigenvector indices for milk production of Holstein cows

The objectives of the present study were to estimate genetic parameters of monthly test-day milk yield (TDMY) of the first lactation of Brazilian Holstein cows using random regression (RR), and to compare the genetic gains for milk production and persistency, derived from RR models, using eigenvector indices and selection indices that did not consider eigenvectors. The data set contained monthly TDMY of 3,543 first lactations of Brazilian Holstein cows calving between 1994 and 2011. The RR model included the fixed effect of the contemporary group (herd-month-year of test days), the covariate calving age (linear and quadratic effects), and a fourth-order regression on Legendre orthogonal polynomials of days in milk (DIM) to model the population-based mean curve. Additive genetic and nongenetic animal effects were fit as RR with 4 classes of residual variance random effect. Eigenvector indices based on the additive genetic RR covariance matrix were used to evaluate the genetic gains of milk yield and persistency compared with the traditional selection index (selection index based on breeding values of milk yield until 305 DIM). The heritability estimates for monthly TDMY ranged from 0.12 ± 0.04 to 0.31 ± 0.04. The estimates of additive genetic and nongenetic animal effects correlation were close to 1 at adjacent monthly TDMY, with a tendency to diminish as the time between DIM classes increased. The first eigenvector was related to the increase of the genetic response of the milk yield and the second eigenvector was related to the increase of the genetic gains of the persistency but it contributed to decrease the genetic gains for total milk yield. Therefore, using this eigenvector to improve persistency will not contribute to change the shape of genetic curve pattern. If the breeding goal is to improve milk production and persistency, complete sequential eigenvector indices (selection indices composite with all eigenvectors) could be used with higher economic values for persistency. However, if the breeding goal is to improve only milk yield, the traditional selection index is indicated.     

Estimated heterogeneity of phenotypic variance of test-day yield with a structural variance model

First-lactation test-day milk, fat, and protein yields from New York, Wisconsin, and California herds from 1990 through 2000 were adjusted additively for age and lactation stage. A random regression model with third-order Legendre polynomials for permanent environmental and genetic effects was used. The model included a random effect with the same polynomial regressions for 2 yr of calvings within herd (herd-time effect) to provide herd-specific lactation curves that can change every 2 yr. (Co)variance components were estimated using expectation-maximization REML simultaneously with phenotypic variances that were modeled using a structural variance model. Maximum heritability for test-day milk yield was estimated to be approximately 20% around 200 to 250 d in milk; heritabilities were slightly lower for test-day fat and protein yields. Herd-time effects explained 12 to 20% of phenotypic variance and had the greatest impact at start of lactation. Variances of test-day yields increased with time, subclass size, and milking frequency. Test month had limited influence on variance. Variance increased for cows in herds with low and high milk yields and for early and late lactation stages. Repeatabilities of variances observed for a given class of herd, test-day, and milking frequency were 14 to 17% across nested variance subclasses based on lactation stage.