Survival rates and productive herd life of dairy cattle in the United States

Survival rates and productive herd life were examined for 13.8 million US dairy cows that calved from January 1, 1980, through March 2, 2005. Cows that left the herd for dairy purposes or were from herds that discontinued Dairy Herd Improvement testing were excluded from any calculations to prevent underestimation of population longevity. Mean lactation length for cows without subsequently recorded lactations ranged from 205 to 235 d across breed-parity subsets and were 4 to 29 d longer for parities 2 through 7 than for parity 1. Mean survival rates were 73% to parity 2; 50% to parity 3; 32% to parity 4; and 19, 10, 5, and 2% to parities 5 through 8, respectively. The mean number of parities for Holsteins declined from 3.2 for those first calving in 1980 to 2.8 for those first calving in 1994. Mean numbers of parities for other breeds first calving in 1994 were 2.9 for Ayrshires and Brown Swiss, 2.4 for Guernseys, and 3.2 for Jerseys. Breed means for productive herd life (through parity 8) ranged from 28 to 36 mo. All regressions of mean number of parities or mean productive herd life on year were negative. The trend for decline of many of those indicators of longevity slowed or ended after the early 1990s. Between 31 (Jersey) and 39% (Guernsey) of herds were made up of first-calf heifers.     

Analysis of the relationship between somatic cell score and functional longevity in Canadian dairy cattle

The aim of this study was to assess the level of somatic cell count (SCC) and to explore the impact of somatic cell score (SCS) on the functional longevity of Canadian dairy cattle by using a Weibull proportional hazards model. Data consisted of 1,911,428 cows from 15,970 herds sired by 7,826 sires for Holsteins, 80,977 cows in 2,036 herds from 1,153 sires for Ayrshires, and 53,114 cows in 1,372 herds from 1,758 sires for Jerseys. Functional longevity was defined as the number of days from the first calving to culling, death, or censoring. The test-day SCC was transformed to a linear score, and the resulting SCS were averaged within each lactation. The average SCS were grouped into 10 classes. The statistical model included the effects of stage of lactation; season of production; annual change in herd size; type of milk recording supervision; age at first calving; effects of milk, fat, and protein yields, calculated as within-herd-year-parity deviations; herd-year-season of calving; SCS class; and sire. The relative culling rate was calculated for animals in each SCS class after accounting for the aforementioned effects. The overall average SCC for Holsteins was 167,000 cells/mL, for Ayrshires was 155,000 cells/mL, and for the Jerseys was 212,000 cells/mL. In all breeds there were no appreciable differences in the relative risk of culling among classes of SCS breed averages (i.e., up to a SCS of 5). However, as the SCS increased beyond the breed average, the relative risk of cows being culled increased considerably. For instance, Holstein, Ayrshire, and Jersey cows with the highest classes of SCS had, respectively, a 4.95, 6.73, and 6.62 times greater risk of being culled than cows with average SCS.     

Indirect prediction of herd life in Guernsey dairy cattle

Production and type data were used to investigate the relationships of these traits with herd life data in US Guernsey cows that calved from 1985 through 1990. Two definitions of herd life were used: actual days from birth to disposal (true herd life) and herd life adjusted for milk production (functional herd life). Genetic parameters were calculated with data from cows that had an opportunity to reach 84 mo of age (n = 18,725). Linear type traits were preadjusted for stage of lactation and age at classification. True herd life was preadjusted for age at first calving and for functional herd life, within herd-year quartile ranking for milk yield. The (co)variance components for true and functional herd life, milk, fat, protein, and 15 linear type traits were estimated with multiple-trait REML in an animal model. Heritability estimates for true and functional herd life were 0.12 for both traits. Estimated genetic correlations of herd life with body size traits were from -0.14 to -0.29, with feet and leg traits were from -0.10 to 0.06, and with udder traits were from -0.09 to 0.24. These correlation parameters were used for indirect prediction of herd life from available production and type information in Guernseys.     

Short communication: Modification of genetic evaluation of herd life from a three-trait to a five-trait model in Canadian dairy cattle

The national genetic evaluation of herd life for Canadian dairy breeds was modified from a 3-trait to a 5-trait animal model. The genetic evaluation incorporates information from daughter survival (direct herd life) and information from conformation, fertility, and udder health traits that are related to longevity (indirect herd life). Genetic evaluations for direct herd life were based on cows’ survival from first calving to 120 days in milk (DIM), from 120 to 240 DIM, from 240 DIM to second calving, survival to third calving, and survival to fourth calving, which were analyzed using a multiple-trait animal model. Sire evaluations obtained for each of the 5 survival traits were combined into an overall sire evaluation for direct herd life. Sire evaluations for indirect herd life were based on an index of sire evaluations for dairy strength, feet and legs, overall mammary, rump angle, somatic cell score, milking speed, nonreturn rate in cows, and interval from calving to first service. A multiple-trait sire model based on multiple-trait across-country evaluation methodology was used to combine direct and indirect genetic evaluations for herd life into an overall genetic evaluation for herd life. Sire evaluations for herd life were expressed as an estimated transmitting ability for the number of lactations. The transmitting ability represents expected differences among daughters for herd life; and the average herd life was set to 3 lactations.     

Genetic and environmental analysis of female calf survival in the Israel Holstein cattle population

The objectives were to estimate the effects of various environmental factors on female calf survival of Israeli Holsteins, to estimate the economic value of calf survival under Israeli conditions, to estimate the genetic and environmental variance components for calf and cow survival using the individual animal model, to perform GWAS analyses of survival to first calving and herd life after first calving, to estimate the genetic and environmental trends for calf survival since 1985, to estimate genetic correlations of calf survival with the traits included in the current Israeli breeding index, and to estimate the consequences of inclusion of calf survival in the national selection index. Mean calf survival rate of Israeli Holsteins from 2001 through 2008 was 0.85, and the mean economic value of survival to first calving was $526. Birth month, gestation length, dystocia, and twin birth significantly affected calf survival rate. Dystocia and twin birth each reduced survival rate by 0.034. Survival rate was highest for calves born in October and lowest for calves born in February. The difference between these months was 3.4%. Maximum survival was at a gestation length of 276 d, the mean gestation length for this population. Survival rate was reduced to 0.76 for calves born after a gestation length of 260 d. The individual animal model was applied for all the genetic analyses. Heritability for calf survival to first calving, as estimated by REML, was 0.009, whereas heritability of herd life from first calving was 0.15. The complete data set for genetic analysis of survival to first calving included 1,235,815 calves born between 1985 and 2017. Annual genetic and phenotypic trends for calf survival were 0.019 and 0.015%, respectively. Correlations of transmitting abilities of 226 sires born since 2010 for calf survival with the traits included in the Israeli breeding index were significant only for the maternal effects of dystocia and stillbirth. The GWAS analysis was based on the transmitting abilities of 1,493 bulls with genotypes and reliabilities >0.5 for calf survival and cow herd life. There were 7 single nucleotide polymorphisms with coefficients of determination >0.03 for calf survival and 12 single nucleotide polymorphisms with coefficients of determination >0.05 for cow survival. There was no overlap between the genome-wide significant markers for the GWAS analyses of calf survival and cow herd life. This corresponds to the conclusion from the REML results and the low correlations between the sire evaluations that the genetic control of the 2 traits are not similar. Inclusion of calf survival in the Israeli breeding would result in a 0.5% increase in calf survival over 10 yr but reduce progress for the other traits by 8%.     

Does clinical mastitis in the first 100 days of lactation 1 predict increased mastitis occurrence and shorter herd life in dairy cows?

The objectives of this study were to estimate the direct effects of clinical mastitis (CM) occurring in early productive life (defined as the first 100 d of the first lactation) of Holstein dairy cows on the future rate of CM occurrence and on the length of total productive lifetime. Information on CM cases and other data occurring in 55,144 lactations in 24,831 cows in 5 New York State Holstein herds was collected from January 2004 until February 2014. For the first objective, a generalized linear mixed model with a Poisson distribution was used to study the effects of CM cases occurring in the first 100 d of a cow's first lactation, as well as farm indicator and number of days in the cow's lifetime, on the future lifetime rate of CM. Only cows that had completed their productive life [i.e., all had been culled (or sold) or had died; n = 14,440 cows] were included in this analysis. For the second objective, a Cox proportional hazards model was used to study the effects of CM cases occurring in the first 100 d of a cow's first lactation on the length of total productive lifetime. The model was stratified by farm. All 24,831 cows were included in this analysis with right censoring. Cows experienced between 0 and 4 CM cases in the first 100 d of lactation 1. Over their lifetime, cows experienced between 0 and 25 CM cases. During the study period, 10% of all cows died and nearly half of all cows were culled. The average length of productive life, including censored observations, was 2.0 yr after first calving. Compared with cows having no CM cases in the first 100 d of lactation 1, cows with 1 CM case in that time period had a 1.5 times higher rate of total number of CM cases over their lifetime. Cows with 2 (or 3 or more) CM cases in the first 100 d of lactation 1 had a 1.7 times (or 2.6 times) higher rate of total number of CM cases over their lifetime. For each additional CM case occurring in the first 100 d of lactation 1, the hazard rate of culling increased by 34%. Given economic conditions for preferentially culling mastitic cows, the study findings may help farmers make optimal decisions with regard to culling of such cows.     

A stochastic model simulating pathogen-specific mastitis control in a dairy herd

The objective of this study was to develop a model simulating mastitis control in dairy herds and to investigate how sensitive the model is when varying the effect parameters according to the uncertainty. The model simulates 9 pathogen-specific mastitis types, each of which can be subclinical or clinical. The clinical cases can be 1 of 4 severities defined according to the effect of the mastitis case: mild, moderate, severe, and permanent effect. The risk factors include lactation stage, parity, yield level, previous diseases, season, and contagious spread of the infection from herd mates. Occurrence of mastitis is modeled to have direct effects on feed intake, body weight, milk yield, somatic cell count in the milk, subsequent mastitis cases within the cow and in herd mates, voluntary and involuntary culling, mortality, and milk withdrawal. Thirty-five scenarios were simulated to study model behavior and model sensitivity. The consequences per cow/yr of mastitis in the default simulated herd included 0.42 clinical mastitis occurrences, 0.56 subclinical mastitis occurrences, loss of 385-kg milk yield, a 1.3% reduced feed intake, 61-kg milk withdrawal and €146 in reduced economic net return. Based on scenarios demonstrating model behavior and sensitivity analysis, the model appears to produce valid consequences of mastitis control strategies. Representation of the effect of subclinical mastitis and of variation in mastitis severity was concluded in this study to be important when modeling mastitis economics in a dairy herd. The model offers the opportunity to study the long-term herd specific effects of a wide range of control strategies against mastitis.     

Comparison between a Weibull proportional hazards model and a linear model for predicting the genetic merit of US Jersey sires for daughter longevity

Predicted transmitting abilities (PTA) of US Jersey sires for daughter longevity were calculated using a Weibull proportional hazards sire model and compared with predictions from a conventional linear animal model. Culling data from 268,008 Jersey cows with first calving from 1981 to 2000 were used. The proportional hazards model included time-dependent effects of herd-year-season contemporary group and parity by stage of lactation interaction, as well as time-independent effects of sire and age at first calving. Sire variances and parameters of the Weibull distribution were estimated, providing heritability estimates of 4.7% on the log scale and 18.0% on the original scale. The PTA of each sire was expressed as the expected risk of culling relative to daughters of an average sire. Risk ratios (RR) ranged from 0.7 to 1.3, indicating that the risk of culling for daughters of the best sires was 30% lower than for daughters of average sires and nearly 50% lower than than for daughters of the poorest sires. Sire PTA from the proportional hazards model were compared with PTA from a linear model similar to that used for routine national genetic evaluation of length of productive life (PL) using cross-validation in independent samples of herds. Models were compared using logistic regression of daughters' stayability to second, third, fourth, or fifth lactation on their sires' PTA values, with alternative approaches for weighting the contribution of each sire. Models were also compared using logistic regression of daughters' stayability to 36, 48, 60, 72, and 84 mo of life. The proportional hazards model generally yielded more accurate predictions according to these criteria, but differences in predictive ability between methods were smaller when using a Kullback-Leibler distance than with other approaches. Results of this study suggest that survival analysis methodology may provide more accurate predictions of genetic merit for longevity than conventional linear models.     

Using conformation traits to improve reliability of genetic evaluation for herd life based on survival analysis

Genetic evaluation for herd life based on survival analysis utilizes information available on all animals, dead (uncensored) and alive (censored), but the reliability of bulls' breeding values depends only on the number of uncensored daughters. Therefore, information on correlated conformation traits scored on daughters during their first lactation may be essential for the evaluation of young bulls with mostly censored daughters. Currently available programs for genetic evaluation based on survival analysis cannot combine indirect information on conformation traits with direct information on herd life, nor can they estimate genetic covariances between herd life and conformation traits. In this study, an alternative approach has been developed and tested using data on Swiss Simmental and Red & White cattle. Genetic covariances were approximated using breeding values for herd life from a survival analysis and BLUP breeding values for 26 linear conformation traits from a separate multivariate analysis. An index combining direct breeding values for herd life and indirect breeding values obtained from conformation traits was constructed. The relative weighting of both information sources varied depending on the amount of available information. The maximum reliability based only on conformation traits was 0.64. Except for old bulls with >100 uncensored daughters, the combined reliability was always higher than the direct reliability from survival analysis.     

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.     

Impact of calving ease on functional longevity and herd amortization costs in Basque Holsteins using survival analysis

The aim of this study was to analyze the impact of calving ease (CE) on functional longevity of Basque Holsteins, using a Weibull proportional hazards model. The data considered for the analysis were 53,353 calving records from 25,810 Holstein cows distributed across 781 herds and sired by 746 bulls. The effects included in the statistical model were age at first calving, stage of lactation, interaction between year and season of calving, 305-d adjusted milk yield, CE, herd, and sire. Calving ease was considered as a time-dependent covariate and, as was the case for the rest of covariates included in the model, had a significant effect on functional longevity. Calvings needing assistance or surgery increased culling risk by 18%, when compared with unassisted calvings. The effect of CE on length of productive life in primiparous and multiparous cows was also investigated. A second analysis was performed replacing the CE effect with the interaction between parity and CE to evaluate the effect of CE in primiparous and multiparous cows. An increase in calving difficulty had a greater impact on culling during first lactations than in subsequent ones. Therefore, difficult calvings, mainly at first parities, had a high impact on herd amortization costs, increasing them by 10% in relation to easy calvings. Therefore, calving difficulty should be avoided as much as possible, especially in primiparous cows, to avoid reduction of profitability.     

Analysis of the relationship between type traits and functional survival in US Holstein cattle using a Weibull proportional hazards model

Survival analysis with a Weibull proportional hazards model was used to evaluate the effects of 15 linear type traits, 5 composite traits, and final score on the functional longevity of US Holstein cows. Culling data and type classification scores (measured in first lactation) from 891,524 cows with first calving from 1993 to 2000 were used. The data were divided into 9 geographical regions to determine whether the relationship between type traits and longevity differed according to climate or management system. Functional survival was defined as the number of days from first calving until culling or censoring, after correction for 305-d mature equivalent combined fat and protein yield. The Weibull model included time-dependent effects of herd-year-season, parity-stage of lactation, and within herd-year quintile ranking for combined fat and protein yield (nested within biennium), as well as time-independent effects of age at first calving and type classification score (type traits were analyzed one at a time). Type classification scores were rounded to the nearest 5 points, and the impact of each type trait on functional survival in each region was evaluated. Mean failure time ranged from 694 d in the South to 758 d in the North East. Risk of culling differed by region for several linear type traits, and differences were greatest for regions that were most dissimilar in climate and herd management (e.g., South East, East North Central, and West). Udder depth, fore udder attachment, udder cleft, and rear legs side view were consistently associated with functional longevity, regardless of region, but, the importance of some secondary traits, such as stature or dairy form, differed by region. The survival model applied in this study easily described both linear and nonlinear relationships between type traits and longevity while accounting for important time-dependent and time-independent explanatory variables.     

Analysis of the relationship between type traits, inbreeding, and functional survival in Jersey cattle using a Weibull proportional hazards model

A Weibull proportional hazards model was used to analyze the effects of 13 linear type traits, final score, and inbreeding on the functional survival of 268,008 US Jersey cows in 2416 herds with first calving from 1981 to 2000. Functional survival was defined as the number of days from first calving until involuntary culling or censoring. The statistical model included the time-dependent effects of herd-year-season of calving, parity by stage of lactation interaction, and within-herd-year quintile for mature equivalent milk yield, as well as the time-independent effects of inbreeding, age at first calving, and linear type traits or final score (analyzed one at a time). Each type trait was divided into 10 classes, and the relative risk of involuntary culling was calculated for animals in each class after accounting for the aforementioned management factors. Type traits with the greatest contribution to the likelihood function were udder depth, fore udder attachment, front teat placement, and udder support. Cows with low scores for these traits had a risk of culling that was 1.3 to 1.8 times that of cows with intermediate scores. Cows with high scores for udder depth and udder support had a risk of culling only 0.7 to 0.85 as great as that of cows with intermediate scores. Intermediate scores were desirable for rear leg set, dairy form, and strength, but stature, rump angle, and rump width had negligible effects on survival. Cows with low final scores had a risk of culling that was 1.35 times that of cows with intermediate scores, whereas cows with high final scores had a risk of culling that was 0.8 times that of cows with intermediate scores. Animals with inbreeding coefficients greater than 10% had a slightly higher risk of culling than animals with inbreeding coefficients less than 5%.     

Genetic analysis of herd life in Quebec Holsteins using Weibull models

Survival analysis methodologies were used to study herd life in Canadian Holstein cows. Herd life was defined as true herd life or the length of time between first calving and censoring. True herd life adjusted for 305-d milk production was defined as functional herd life. Lifetime record (censored or completed) were from 331,147 Holstein cows registered in the Programme d'Analyse des Troupeaux Laitiers du Québec (PATLQ) that calved for the first time between March 1, 1981 and March 31, 1995. The Weibull (proportional hazards) model used to analyze true herd life and functional herd life contained a Weibull baseline hazard function and the time-dependent effects of year of first calving, lactation number by stage of lactation, annual change in herd size and herd-year (random), and the time-independent effects of the milk recording option (supervised or not) and age at first calving. The model for functional herd life included also the time-dependent effect of herd-year-parity class of 305-d milk production. Genetic differences between sires with regard to the hazard function of their daughters was clearly demonstrated. The hazard rate followed a different pattern in later lactations, particularly in the first 240 d in milk. Older age at first calving was found to be associated with higher risks of culling. Changes in herd size had a small impact on the hazard function of animals. The hazard decreased as production of the cow increased. Heritability in the log scale was 0.09 for true herd life and 0.08 for functional herd life, but when heritability was expressed on the original scale, the estimates for the two traits were 0.19 and 0.15, respectively. The difference in the median survival between a bull with an estimated transmitting ability of 0.6 and another bull with an estimated transmitting ability of 1.3 was 690 d or 1.7 lactations. Rank correlations between the official estimated transmitting abilities for true herd life and functional herd life and those obtained in this study were 0.62 and 0.66, respectively.     

Genetic evaluation of fertility traits of dairy cattle using a multiple-trait animal model

A genetic evaluation system was developed for 5 fertility traits of dairy cattle: interval from first to successful insemination and nonreturn rate to 56 d of heifers, and interval from calving to first insemination, nonreturn rate to 56 d, and interval first to successful insemination of cows. Using the 2 interval traits of cows as components, breeding values for days open were derived. A multiple-trait animal model was applied to evaluate these fertility traits. Fertility traits of later lactations of cows were treated as repeated measurements. Genetic parameters were estimated by REML. Mixed model equations of the genetic evaluation model were solved with preconditioned conjugate gradients or the Gauss-Seidel algorithm and iteration on data techniques. Reliabilities of estimated breeding values were approximated with a multi-trait effective daughter contribution method. Daughter yield deviations and associated effective daughter contributions were calculated with a multiple trait approach. The genetic evaluation software was applied to the insemination data of dairy cattle breeds in Germany, Austria, and Luxembourg, and it was validated with various statistical methods. Genetic trends were validated. Small heritability estimates were obtained for all the fertility traits, ranging from 1% for nonreturn rate of heifers to 4% for interval calving to first insemination. Genetic and environmental correlations were low to moderate among the traits. Notably, unfavorable genetic trends were obtained in all the fertility traits. Moderate to high correlations were found between daughter yield-deviations and estimated breeding values (EBV) for Holstein bulls. Because of much lower heritabilities of the fertility traits, the correlations of daughter yield deviations with EBV were significantly lower than those from production traits and lower than the correlations from type traits and longevity. Fertility EBV were correlated unfavorably with EBV of milk production traits but favorably with udder health and longevity. Integrating fertility traits into a total merit selection index can halt or reverse the decline of fertility and improve the longevity of dairy cattle.     

Relationship between herd size and measures of animal welfare on dairy cattle farms with freestall housing in Germany

The objective of this study was to examine the association of herd size with animal welfare in dairy cattle herds. Therefore, 80 conventional dairy cattle farms were classified by the number of cows into 4 herd size classes, C1 (<100 cows), C2 (100–299 cows), C3 (300–499 cows), and C4 (≥500 cows), and assessed using multiple animal-based measures of the Welfare Quality Assessment protocol for dairy cattle. Data were recorded from April 2014 to September 2016 by an experienced single assessor in northern Germany. Each farm was visited 2 times at an interval of 6 mo (summer period and winter period) to avoid seasonal effects on the outcome. The average herd size was 383 ± 356 Holstein-Friesian cows (range 45 to 1,629). Only farms with freestall (cubicle) housing and a maximum of 6 h access to pasture per day were included in the study. Data were statistically analyzed using a generalized linear mixed model. None of the farms reached the highest overall rating of “excellent.” The majority of the farms were classified as “enhanced” (30%) or “acceptable” (66%), and at 6 assessments the farms were rated as “not classified” (4%). Regarding single indicators, mean trough length per cow, percentage of cows with nasal discharge, and vulvar discharge increased with increasing herd size, whereas it was vice versa for displacements of cows. Percentage of lean cows, percentage of dirty lower legs, and duration of the process of lying down showed a curvilinear relationship with the number of cows per farm. Herd size was not associated with any other measures of the Welfare Quality protocol. In conclusion, herd size effects were small, and consequently herd size cannot be considered as a feasible indicator of the on-farm animal welfare level. Housing conditions and management practices seem to have a greater effect on animal welfare than the number of dairy cows per farm.     

Genetic parameters derived from using a biological model of lactation on records of commercial dairy cows

The object of this study was to investigate the genetics of lactation curve parameters derived from a biological model of lactation and the relationships among them. This biological model fitted 2 logistic curves to mimic the initial increase in milk secretory cell numbers in early lactation and the progression of apoptosis in late lactation. Records from 82,255 Holstein-Friesian heifers from commercial dairy herds in the United Kingdom, recorded from 1994 to 2003, were analyzed. The heritabilities of 2 lactation curve parameters, maximum secretion potential and relative cell death rate, were 0.27 and 0.08 respectively. Maximum secretion potential was highly genetically correlated with peak yield (0.99), and relative cell death rate was highly correlated with persistency of lactation (0.84). Heritability values for the traits analyzed showed a characteristic pattern. Total milk yield traits, maximum secretion potential, and peak yield had similar and moderate heritabilities (∼0.3). Traits associated with late lactation had lower heritability values (∼0.1), whereas day of peak yield and early lactation traits had little genetic variation. The permanent environmental variance of the various traits ranged from 0.08 to 0.26 of the phenotypic variance. Parameters from the 2 logistic curves were not highly correlated, suggesting that selection programs could be devised to exploit genetic variation in both aspects of lactation independently.