Genetic relationships between weight loss in early lactation and daily milk production throughout lactation in Holstein cows.

After calving, high-yielding dairy cows mobilize body reserves for energy, sometimes to the detriment of health and fertility. This study aimed to estimate the genetic correlation between body weight loss until nadir and daily milk production (MY24) in first- (L1) and second-lactation (L2) Holstein cows. The data set included 859,020 MY24 records and 570,651 daily raw body weight (BWr) phenotypes from 3,989 L1 cows, and 665,361 MY24 records and 449,449 BWr phenotypes from 3,060 L2 cows, recorded on 36 French commercial farms equipped with milking robots that included an automatic weighing platform. To avoid any bias due to change in digestive content, BWr was adjusted for variations in feed intake, estimated from milk production and BWr. Adjusted body weight was denoted BW. The genetic parameters of BW and MY24 in L1 and L2 cows were estimated using a 4-trait random regression model. In this model, the random effects were fitted by second-order Legendre polynomials on a weekly basis from wk 1 to 44. Nadir of BW was found to be earlier than reported in the literature, at 29 d in milk, and BW loss from calving to nadir was also lower than generally assumed, close to 29 kg. To estimate genetic correlations between body weight loss and production, we defined BWL5 as the loss of weight between wk 1 and 5 after calving. Genetic correlations between BWL5 and MY24 ranged from -0.26 to 0.05 in L1 and from -0.11 to 0.10 in L2, according to days in milk. These moderate to low values suggest that it may be possible to select for milk production without increasing early body mobilization.

Association between body condition genomic values and feed intake, milk production, and body weight in French Holstein cows.

Body condition score (BCS) offers a good estimate of the amount of stored fat on the body, and its variations can be used as a proxy for energy balance. Many countries have implemented a genomic evaluation of BCS, including France, where estimated breeding values are based on an individual BCS determination during the first lactation. In this article, we investigate the degree to which this genomic estimated breeding value based on a single phenotype record per cow might reflect different profiles of body reserves throughout lactation and be used to predict, and perhaps limit, their mobilization during early lactation. We also investigate whether selection on BCS affects other traits. A data set including 686 lactations of 435 Holstein cows from 3 experimental farms not used in the reference population for genomic evaluation was used to estimate the effects of the BCS direct genomic value (iBCS) on BCS, body weight, feed intake, milk production, and fat and protein contents throughout the lactation period. For each trait, the model included different iBCS regressions and an effect of the direct genomic value of the trait itself when available. It thus appeared that cows with a positive iBCS always had a higher BCS than negative iBCS cows, whatever the lactation stage, and that this difference increased during the first 6 mo to reach a difference of 0.8 point. A similar effect was seen regarding body weight, but it was the opposite for milk production, with negative iBCS cows producing slightly more milk (difference of about 3% over lactation). Feed intake increased slightly faster at the beginning of lactation for cows with positive iBCS. Therefore, iBCS is a promising tool that could help to limit intense mobilization during early lactation. Should feed efficiency be included in the breeding goal, greater attention should be paid to BCS to avoid further body mobilization in early lactation.

Opportunities for genomic selection of cheese-making traits in Montbéliarde cows.

As part of the From'MIR project, traits related to the composition and cheese-making properties (CMP) of milk were predicted from 6.6 million mid-infrared spectra taken from 410,622 Montbéliarde cows (19,862 with genotypes). Genome-wide association studies of imputed whole-genome sequences highlighted candidate SNPs that were then added to the EuroG10K BeadChip, which is routinely used in genomic selection. In the present study, we (1) assessed the reliability of single-step genomic BLUP breeding values (ssEBVs) for cheese yields, coagulation traits, and casein and calcium content generated from test-day records of the first 3 lactations, (2) estimated realized genetic trends for these traits over the last decade, and (3) simulated different cheese-making breeding objectives and estimated the responses for CMP as well as for other traits currently selected in the Montbéliarde breed. To estimate the reliability of ssEBVs, the available data were split into 2 independent training and validation sets that respectively contained cows with the oldest and the most recent lactation data. The training set included 155,961 cows (12,850 with genotypes) and was used to predict ssEBVs of 2,125 genotyped cows in the validation set. We first tested 4 models that included either lactation (LACT) or test-day (TD) records from the first (1) or the first 3 (3) lactations, giving equal weight to all 50K SNP effects. Mean reliabilities were 61%, 62%, 63%, and 64% for the LACT1, LACT3, TD1, and TD3 models, respectively. Using the most accurate model (TD3), we then compared the reliabilities of 3 scenarios with: SNPs from the Illumina BovineSNP50 BeadChip only, equally weighted (50K); 50K SNPs plus additional candidate SNPs, equally weighted (50K+); and 50K and candidate SNPs with additional weight given to 7 to 14 candidate SNPs, depending on the trait (CAND). The 50K+ and CAND scenarios led to similar mean reliabilities (67%) and both outperformed the 50K scenario (64%), whereas the CAND scenario generated the less biased ssEBVs. To assess genetic trends, SNP effects were estimated with a single-step GBLUP based on the TD3 model and the 50K scenario applied to the whole population (2.6 million performance records from 190,261 cows and 423,348 animals in the pedigree, of which 21,874 were genotyped) and then applied to 50K genotypes of 21,171 males and 311,761 females. We detected a positive genetic trend for all CMP during the last decade, probably due to selection for an increase in milk protein and fat content in Montbéliarde cows. Finally, we compared the selection responses to 3 different breeding objectives: the current Montbéliarde total merit index (TMI) and 2 alternative scenarios that gave a weight of 70% to TMI and the remaining 30% to either milk casein content (TMI-COMP) or a combination of 3 CMP (TMI-Cheese). The TMI-Cheese scenario yielded the best responses for all the CMP analyzed, whereas values in the TMI-COMP scenario were intermediate, with a slight effect on other traits currently included in TMI. Based on these results, a program of genomic evaluation for CMP predicted from mid-infrared spectra was designed and implemented for the Montbéliarde breed.

Exploring simultaneous perturbation profiles in milk yield and body weight reveals a diversity of animal responses and new opportunities to identify resilience proxies.

Livestock husbandry aims to manage the environment in which animals are reared to enable them to express their production potential. However, animals are often confronted with perturbations that affect their performance. Evaluating effects of these perturbations on animal performance could provide metrics to quantify and understand how animals cope with their environment, and therefore to better manage them. Body weight (BW) and milk yield (MY) dynamics over lactation may be used for this purpose. The goal of this study was to estimate an unperturbed performance trajectory using a differential smoothing approach on both MY and BW time series, and then to identify the perturbations and extract their phenotypic features. Daily MY and BW records from 490 primiparous Holstein cows from 33 commercial French herds were used. From the fitting procedure, estimated unperturbed performance trajectories of BW and MY were clustered into 3 groups. After the fitting procedure, 1,754 deviations were detected in the MY time series and 964 were detected in the BW time series across all cows. Overall, 425 of these deviations were detected during the same period (±10 d) in both MY and BW time series, 76 of which started at the same time. Results suggest that combining various individual dynamic measures and revealing the relationship that exists between them could be of great value in obtaining reliable estimates of resilience components in large populations.

Estimation of the effects of selection on French Large White sow and piglet performance during the suckling period.

The effects of 21 yr of selection were estimated for sow and piglet performance during the suckling period in a French Large White (LW) pig population using frozen semen. Two experimental groups (EXP = L77 and L98) were produced by inseminating LW sows with either stored frozen semen from 17 LW boars born in 1977 (EXP = L77) or with fresh semen from 23 LW boars born in 1998 (EXP = L98). Seventy-four L77 and 89 L98 randomly chosen females were mated to 15 L77 and 15 L98, respectively, randomly chosen boars for 6 successive parities. They produced 2,796 L77 progeny (G77) and 3,529 L98 progeny (G98) piglets including stillbirths. To disentangle direct and maternal effects on piglet growth, a 2 × 2 factorial design was set by cross-fostering half-litters across genetic groups the day after farrowing, resulting in mixed G77/G98 litters nursed by either L77 or L98 sows. Piglet traits investigated included individual weight at birth (IWB), at 21 d of age (IW21d), and at weaning at 4 wk of age (IWW) and ADG from birth to 21 d of age (ADG21d) and from birth to weaning (ADGBW) as well as probability of stillbirth, probability of mortality on the first day after farrowing and from d 2 to weaning. Sow traits analyzed included weight before farrowing and at weaning, feed intake, milk production, colostrum, and milk composition. The variability of performance across genetic groups and litters was also investigated. The data were analyzed using generalized (piglet mortality) or linear mixed models (other traits). Results showed an increase in IWB (+240 ± 72 g in 21 yr for IWB adjusted for total number born), and a negative maternal genetic trend was observed on piglet growth during the suckling period (e.g., +33 ± 13 g/d in 21 yr for ADG21d, that is, 14% of the mean), whereas direct genetic effects remained unchanged. Piglets from L98 litters also had a 40% larger probability of being stillborn and a 28% larger probability of dying on d 1 and had a more heterogeneous IWB (358 vs. 336 g; < 0.001) and growth during the suckling period (60 vs. 56 g/d; < 0.001). Sows from L77 and L98 experimental groups did not differ in weight, feed intake, colostrum, and milk composition. These results give evidence of negative correlated effects of selection for piglet traits related to robustness. These adverse effects are at least partly of maternal origin.

Estimation of the effects of selection on French Large White reproductive performance using frozen semen.

Genetic trends for male and female sexual development and components of litter size and weight at birth (LB) as well as traits related to sow lifetime productivity were estimated in a French Large White (LW) pig population using frozen semen. Two experimental groups (EXP = L77 and L98) were produced by inseminating LW sows with either stored frozen semen from 17 LW boars born in 1977 (EXP = L77) or with semen from 23 LW boars born in 1998 (EXP = L98). In each group, about 100 males were measured for testicular development and bulbo-urethral gland development, and 90 gilts were checked for puberty, among which about 50 gilts were measured for their first ovulation rate at puberty (ORP). The females were then mated to 15 randomly chosen males for each group for 6 successive parities and measured for total number born (TNB), number born alive (NBA), litter weight at birth (LWB), and interval from weaning to first estrus as well as ovulation rate at fertilization (ORF = ovulation rate of gestating sows) on a subset of litters. The data were analyzed using mixed linear models including the fixed effects of EXP, of finishing (sexual development traits) or farrowing group, and parity (LB traits); the random effects of birth litter (sexual development traits) or sow (LB traits); and, when relevant, age or BW as covariates. The homogeneity of residual variances across EXP was also tested. For each trait, the genetic trend was estimated as twice the difference between L98 and L77. Limited differences were observed for sexual development traits, except for ORF, which increased by 0.17 ± 0.06/yr. The residual SD of NBA (2.7 vs. 2.0; = 0.004) and LWB (3.1 vs. 2.5; = 0.03) were larger in L98 sows than in L77 sows. Positive trends were observed for all litter traits (0.22 ± 0.07 ova, 0.12 ± 0.04 piglet, 0.09 ± 0.04 piglet, and 0.23 ± 0.06 kg/yr for ORF, TNB, NBA, and LWB, respectively) but at the expense of a degradation of prenatal and farrowing survival. A negative trend was also observed for length of productive life (-8.9 ± 3.9 d/yr). These results suggest that gains in productivity were associated with a loss of robustness.

Genetics of residual feed intake in growing pigs: Relationships with production traits, and nitrogen and phosphorus excretion traits.

Residual feed intake (RFI) is defined as the difference between the observed ADFI and the ADFI predicted from production and maintenance requirements. The objectives of this study were to evaluate RFI as a selection criterion to improve feed efficiency and its potential to reduce N and P excretion in 4 pig breeds. Data were collected between 2000 and 2009 in French central test stations for 2 dam breeds [French Landrace (LR) and Large White (LWD)], and 2 sire breeds [Large White (LWS) and Piétrain (PP)]. Numbers of recorded pigs were 6407, 10,694, 2342, and 2448 for the LR, LWD, LWS, and PP breeds, respectively. All PP animals were genotyped for the halothane mutation. This data set was used to calculate RFI equations for each of the 4 breeds, and to estimate genetic parameters for RFI together with growth, carcass, and meat quality traits, and N and P excretion during the test period (35 to 110 kg BW). The RFI explained 20.1% in PP, 26.5% in LWS, 27.6% in LWD, and 29.5% in LR of the phenotypic variability of ADFI. The PP breed differed from the others in this respect, probably due to a lower impact of the variation of body composition on ADFI. Heritability estimates of RFI ranged from 0.21 ± 0.03 (LWD) to 0.33 ± 0.06 (PP) depending on the breed. Heritabilities of N and P excretion traits ranged from 0.29 ± 0.06 to 0.40 ± 0.06. The RFI showed positive genetic correlations with feed conversion ratio (FCR) and excretion traits, these correlations being greater in the sire breeds (from 0.57 to 0.86) than in the dam breeds (from 0.38 to 0.53). Compared with FCR, RFI had weaker genetic correlations with carcass composition, growth rate, and excretion traits. Estimates of genetic correlations between FCR and excretion traits were very close to 1 for all breeds. Finally, excretion traits were, at the genetic level, correlated positively with ADFI, negatively with growth rate and carcass leanness, whereas the halothane n mutation in PP was shown to reduce N and P excretion levels. To conclude, new selection indexes including RFI can be envisaged to efficiently disentangle the responses to selection on growth rate and body composition from those on feed efficiency, with favorable impacts on N and P excretions, particularly in sire pig breeds. However, the switch from FCR to RFI in selection indexes should not resolve the genetic antagonism between feed efficiency and meat quality.

Phenotypic prediction based on metabolomic data for growing pigs from three main European breeds.

Predicting phenotypes is a statistical and biotechnical challenge, both in medicine (predicting an illness) and animal breeding (predicting the carcass economical value on a young living animal). High-throughput fine phenotyping is possible using metabolomics, which describes the global metabolic status of an individual, and is the closest to the terminal phenotype. The purpose of this work was to quantify the prediction power of metabolomic profiles for commonly used production phenotypes from a single blood sample from growing pigs. Several statistical approaches were investigated and compared on the basis of cross validation: raw data vs. signal preprocessing (wavelet transformation), with a single-feature selection method. The best results in terms of prediction accuracy were obtained when data were preprocessed using wavelet transformations on the Daubechies basis. The phenotypes related to meat quality were not well predicted because the blood sample was taken some time before slaughter, and slaughter is known to have a strong influence on these traits. By contrast, phenotypes of potential economic interest (e.g., lean meat percentage and ADFI) were well predicted (R(2) = 0.7; P < 0.0001) using metabolomic data.

Efficiency of genomic selection in a purebred pig male line.

Stochastic simulation was used to compare the efficiency of 3 pig breeding schemes based on either traditional genetic evaluation or genomic evaluation. The simulated population contained 1,050 female and 50 male breeding animals. It was selected for 10 yr for a synthetic breeding goal that included 2 traits with equal economic weights and heritabilities of 0.2 or 0.4. The reference breeding scheme, named BLUP-AM, was based on the phenotyping of all candidates (13,770 animals/yr) for Trait 1 and of relatives from 10% of the litters (270 animals/yr) for Trait 2 and on BLUP-Animal Model genetic evaluations. Under the first alternative scenario, named GE-1TP, selection was based on genomic breeding values (GBV) estimated with one training population (TP) made up of candidate relatives phenotyped for both traits, with a size increasing from 1,000 to 3,430 over time. Under the second alternative scenario, named GE-2TP, the GBV for Trait 2 were estimated using a TP identical to that of GE-1TP, but the GBV for Trait 1 were estimated using a large TP made up of candidates that increased in number from 13,770 to 55,080 over time. Over the simulated period, both genomic breeding schemes generated 39 to 58% more accurate EBV for Trait 2 than the reference scheme, resulting in 78 to 128% (GE-1TP) and 63 to 84% (GE-2TP) greater average annual genetic trends for this trait. For Trait 1, GE-1TP was 18 to 24% less accurate than BLUP-AM, reducing average annual genetic trends by 27 to 44%. By contrast, GE-2TP generated 35 to 43% more accurate EBV and 8 to 22% greater average annual genetic trends for Trait 1 than the reference scheme. Consequently, GE-2TP was 27 to 33% more efficient in improving the global breeding goal than BLUP-AM whereas GE-1TP was globally as efficient as the reference scheme. Both genomic schemes reduced the inbreeding rate, the greatest decrease being observed for GE-2TP (-49 to -60% compared with BLUP-AM). In conclusion, genomic selection could substantially and durably improve the efficiency of pig breeding schemes in terms of reliability, genetic trends, and inbreeding rate without any need to modify their current structure. Even though it only generates a small TP, limited annual phenotyping capacity for traits currently only recorded on relatives would not be prohibitive. A large TP is, however, required to outperform the current schemes for traits recorded on the candidates in the latter.

Estimation of genetic trends in French Large White pigs from 1977 to 1998 for growth and carcass traits using frozen semen.

Genetic trends for growth, feed efficiency, composition, and morphometry of carcasses were estimated in a French Large White (LW) pig population using frozen semen. Two groups of pigs were produced by inseminating LW sows with either stored, frozen semen from 17 LW boars born in 1977 or with semen from 23 LW boars born in 1998. In each group, 15 males and 90 females were randomly chosen and mated to produce approximately 1,000 pigs/group. These pigs were performance tested with individual ADFI and serial BW and backfat thickness measurements, slaughtered at 105 kg of BW, and measured for carcass traits. The data were analyzed using mixed linear animal models, including the fixed effect of the experimental group (offspring of 1977 or 1998 boars), the random effect of the additive genetic value of each animal, and, when significant, the fixed effects of sex, fattening batch, and slaughterhouse, the linear regression on BW, and the random effect of the common environment of birth litter. For each trait, the genetic trend was estimated as twice the difference between the 2 experimental groups. Results showed moderately favorable trends for on-test ADG (3.7 +/- 1.3 g/d per year) and feed conversion ratio (-0.014 +/- 0.005 kg/kg per year) in spite of a tendency toward an increase in ADFI (7.6 +/- 4.7 g/yr). A strong reduction in carcass fatness (-0.35 +/- 0.07 mm/yr for carcass average backfat thickness) and a large improvement in carcass leanness (0.31 +/- 0.10 mm(2)/yr and 0.41 +/- 0.08%/yr for loin eye area and carcass muscle content, respectively) were observed. Carcass shape measurements (back and leg length, back width, muscle thickness of hind limbs) were not affected by selection. Serial measurements of BW and backfat thickness showed that the major part of the genetic gains occurred during late growth and that the reduction in the backfat layer was more pronounced in the rear than in the front part of the carcass. The use of frozen semen appears to be a powerful practice to thoroughly investigate changes attributable to selection.

Estimation of genetic trends from 1977 to 2000 for stress-responsive systems in French Large White and Landrace pig populations using frozen semen.

An experimental design aiming at analysing the consequences of genetic selection from 1977 to 1998-2000 on the evolution of stress-responsive systems in the French Large White (LW) and Landrace (LR) pig populations was conducted by INRA and IFIP-Institut du Porc. Large White sows were inseminated with semen from LW boars born in 1977 (frozen semen) or in 1998 and their second-generation offspring were station-tested. Landrace sows were inseminated with semen from LR boars born in 1977 (frozen semen) or in 1999 to 2000, and their progeny was station-tested. Urinary concentration of stress hormones (cortisol and catecholamines) and traits related to carcass composition (estimated carcass lean content (ECLC) and global adiposity) and meat quality (pH 24 h) were measured. For the two populations, selection carried out since 1977 led to an increase in ECLC and a decrease in carcass adiposity. Between 1977 and 1998 to 2000, urinary concentrations of stress hormones were unchanged in the LR breed, but were decreased in the LW breed. Moreover, for the animals generated from LW boars born in 1977 and in 1998, urinary cortisol levels were negatively correlated with ECLC. Therefore, in the LW breed, selection carried out for higher ECLC resulted in a decrease in cortisol production, as well as a reduction of catecholamine production that may be responsible for the lower ultimate pH of meat. Therefore, selection carried out for increased carcass lean content led, in this breed, to large modifications in the functioning of the stress-responsive systems, thereby influencing a large range of physiological regulations and technical properties such as carcass composition and meat pH, which remained however in the normal range for acceptable meat quality.

Optimized management of genetic variability in selected pig populations.

Controlling the increase of coancestry and inbreeding coefficients in selected populations is made possible through calculation of the optimal contributions allowed to breeding animals, given the current situation with regard to genetic diversity, and further, through optimal design of matings. The potential of such an approach for pig breeding was tested by retrospective optimization on the French Landrace population in reference to the matings actually carried out during a 21-week test period. The major constraint was that the average overall estimated breeding value (EBV) should be the same as the observed one, for not decreasing short-term genetic gain. Optimizing breeding allocations to boars would have led one to decrease coancestry and inbreeding coefficients by approximately 20%. This decrease would have even increased to approximately 30%, would have replacements and disposals been optimized after accounting for genetic variability, keeping the same constraint of genetic level identical to the observed one. These results showed the potential value, in the future, of completing each periodical calculation of EBVs by optimizations considering genetic variability and of releasing corresponding information to breeders, in order to enhance maintenance of genetic variability.

Estimation of genetic trends from 1977 to 1998 for farrowing characteristics in the French Large White breed using frozen semen.

The objective of the study was to estimate genetic trends from 1977 to 1998 in the French Large White (LW) breed for stillbirth and associated traits measured at farrowing using frozen semen. Two groups of pigs (G77 and G98) were obtained by inseminating LW sows with semen from LW boars born either in 1977 or in 1998. A second generation was produced by inter se mating in each group. Farrowing was thoroughly supervised through both direct observations and video recording all long farrowing on a total of 137 first- and second-parity litters produced by sows from this second generation (68 G77 and 69 G98 litters, respectively). Measurements included birth time, weight and birth characteristics (including orientation, presence of cyanosis or oedema, membrane obstruction, umbilical cord length/content) of each piglet, as well as sow traits (weight and backfat thickness, farrowing duration, litter size and within-litter variation of weights at birth). The data were analysed using linear or generalised linear mixed models, according to the definition of the trait (continuous or binary data). The importance of several effects to piglet probability of stillbirth was then quantified by computing the reduction of variance associated with the addition of each effect in the model. Litter size did not significantly differ in first parity, but was higher in G98 second-parity sows: the differences for global (including pre partum dead piglets) and total numbers of piglets born per litter were +2.3 ± 1.1 and +1.3 ± 0.6, respectively. G98 sows also had a higher number of stillbirths in both parities (+0.7 ± 0.3 stillborn per litter). Piglets from G98 litters were heavier at birth (+130 ± 40 g for birth weight adjusted for litter size), without any increase in within-litter heterogeneity of birth weight. No significant difference was detected between G77 and G88 groups for farrowing length and the distribution of time interval between piglet births. G98 stillborn piglets had longer and more often empty umbilical cords at birth. G98 piglets born alive also had more often umbilical nodes than G77 piglets. These characteristics were considered as indicators of increased farrowing difficulties and risk of hypoxia at birth in G98 pigs. Time of birth of each piglet, sow fatness at farrowing and time of first placenta expulsion were the main factors of variation of the piglet's probability of stillbirth.

Estimation of genetic trends from 1977 to 1998 of body composition and physiological state of Large White pigs at birth.

Genetic trends for body composition and blood plasma parameters of newborn piglets were estimated through the comparison of two groups of pigs (G77 and G98, respectively) produced by inseminating Large White (LW) sows with semen from LW boars born either in 1977 or in 1998. Random samples of 18 G77 and 19 G98 newborn piglets were used for whole carcass and tissue sampling. Plasma concentrations of glucose, albumin and IGF-1 were determined on 75 G77 and 90 G98 piglets from 18 litters. The G98 piglets had less carcass dry matter, protein and energy (P < 0.01) than their G77 counterparts. When expressed in g/kg birth weight, livers were lighter (P < 0.001) and contained less glycogen (P < 0.01) in G98 piglets, with no difference in the activity of the hepatic glucose-6-phosphatase between G98 and G77 piglets. Concentrations of protein, DNA, RNA in longissimus dorsi muscle were unaffected by selection. Plasma concentrations of glucose (P < 0.05) and IGF-1 (P < 0.01) were lower in G98 than in G77 piglets. On the whole, the results suggest that the improvement in lean growth rate and in sow prolificacy from 1977 to 1998 has resulted in a lower maturity of piglets at birth.