Quantitative trait locus mapping in an F2 Duroc x Pietrain resource population: II. Carcass and meat quality traits

Pigs from the F(2) generation of a Duroc x Pietrain resource population were evaluated to discover QTL affecting carcass composition and meat quality traits. Carcass composition phenotypes included primal cut weights, skeletal characteristics, backfat thickness, and LM area. Meat quality data included LM pH, temperature, objective and subjective color information, marbling and firmness scores, and drip loss. Additionally, chops were analyzed for moisture, protein, and fat composition as well as cook yield and Warner-Bratzler shear force measurements. Palatability of chops was determined by a trained sensory panel. A total of 510 F(2) animals were genotyped for 124 microsatellite markers evenly spaced across the genome. Data were analyzed with line cross, least squares regression interval, mapping methods using sex and litter as fixed effects and carcass weight or slaughter age as covariates. Significance thresholds of the F-statistic for single QTL with additive, dominance, or imprinted effects were determined on chromosome- and genome-wise levels by permutation tests. A total of 94 QTL for 35 of the 38 traits analyzed were found to be significant at the 5% chromosome-wise level. Of these 94 QTL, 44 were significant at the 1% chromosome-wise, 28 of these 44 were also significant at the 5% genome-wise, and 14 of these 28 were also significant at the 1% genome-wise significance thresholds. Putative QTL were discovered for 45-min pH and pH decline from 45 min to 24 h on SSC 3, marbling score and carcass backfat on SSC 6, carcass length and number of ribs on SSC 7, marbling score on SSC 12, and color measurements and tenderness score on SSC 15. These results will facilitate fine mapping efforts to identify genes controlling carcass composition and meat quality traits that can be incorporated into marker-assisted selection programs to accelerate genetic improvement in pig populations.     

Quantitative trait loci mapping in an F2 Duroc x Pietrain resource population: I. Growth traits

Pigs from the F(2) generation of a Duroc x Pietrain resource population were evaluated to discover QTL affecting growth and composition traits. Body weight and ultrasound estimates of 10th-rib backfat, last-rib backfat, and LM area were serially measured throughout development. Estimates of fat-free total lean, total body fat, empty body protein, empty body lipid, and ADG from 10 to 22 wk of age were calculated, and random regression analyses were performed to estimate individual animal phenotypes representing intercept and linear rates of increase in these serial traits. A total of 510 F(2) animals were genotyped for 124 micro-satellite markers evenly spaced across the genome. Data were analyzed with line cross, least squares regression, interval mapping methods using sex and litter as fixed effects. Significance thresholds of the F-statistic for single QTL with additive, dominance, or imprinted effects were determined at the chromosome- and genome-wise levels by permutation tests. A total of 43 QTL for 22 of the 29 measured traits were found to be significant at the 5% chromosome-wise level. Of these 43 QTL, 20 were significant at the 1% chromosome-wise significance threshold, 14 of these 20 were also significant at the 5% genome-wise significance threshold, and 10 of these 14 were also significant at the 1% genome-wise significance threshold. A total of 22 QTL for the animal random regression terms were found to be significant at the 5% chromosome-wise level. Of these 22 QTL, 6 were significant at the 1% chromosome-wise significance threshold, 4 of these 6 were also significant at the 5% genome-wise significance threshold, and 3 of these 4 were also significant at the 1% genome-wise significance threshold. Putative QTL were discovered for 10th-rib and last-rib backfat on SSC 6, body composition traits on SSC 9, backfat and lipid composition traits on SSC 11, 10th-rib backfat and total body fat tissue on SSC 12, and linear regression of last-rib backfat and total body fat tissue on SSC 8. These results will facilitate fine-mapping efforts to identify genes controlling growth and body composition of pigs that can be incorporated into marker-assisted selection programs to accelerate genetic improvement in pig populations.     

Quantitative trait loci for leg weakness traits in a Landrace purebred population

Leg weakness in pigs is a serious problem in the pig industry. We performed a whole genome quantitative trait locus (QTL) analysis to find QTLs affecting leg weakness traits in the Landrace population. Half-sib progeny ( n  = 522) with five sires were measured for leg weakness traits. Whole genome QTL mapping was performed using a half-sib regression-based method using 190 microsatellite markers. No experiment-wide significant QTLs affecting leg weakness traits were detected. However, at the 5% chromosome-wide level, QTLs affecting leg weakness traits were detected on chromosomes 1, 3, 10 and 11 with QTL effects ranging from 0.07 to 0.11 of the phenotypic variance. At the 1% chromosome-wide level, QTLs affecting rear feet score and total leg score were detected on chromosomes 2 and 3 with QTL effects of 0.11 and 0.13 of the phenotypic variance, respectively. On chromosome 3 and 10, some QTLs found in this study were located at nearby positions. The present study is one of the first reports of QTLs affecting fitness related traits such as leg weakness traits, that segregate within the Landrace population. The study also provides useful information for studying QTLs in purebred populations.     

Quantitative trait loci mapping for meat quality and muscle fiber traits in a Japanese wild boar x Large White intercross

Three generations of a swine family produced by crossing a Japanese wild boar and three Large White female pigs were used to map QTL for various production traits. Here we report the results of QTL analyses for skeletal muscle fiber composition and meat quality traits based on phenotypic data of 353 F(2) animals and genotypic data of 225 markers covering almost the entire pig genome for all of the F(2) animals as well as their F(1) parents and F(0) grandparents. The results of a genome scan using least squares regression interval mapping provided evidence that QTL (<1% genome-wise error rate) affected the proportion of the number of type IIA muscle fibers on SSC2, the number of type IIB on SSC14, the relative area (RA) of type I on SSCX, the RA of type IIA on SSC6, the RA of type IIB on SSC6 and SSC14, the Minolta a* values of loin on SSC4 and SSC6, the Minolta b* value of loin on SSC15, and the hematin content of the LM on SSC6. Quantitative trait loci (<5% genome-wise error rate) were found for the number of type I on SSC1, SSC14, and SSCX, for the number of type IIA on SSC14, for the number of type IIB on SSC2, for the RA of type IIA on SSC2, for the Minolta b* value of loin on SSC3, for the pH of loin on SSC15, and for the i.m. fat content on SSC15. Twenty-four QTL were detected for 11 traits at the 5% genome-wise level. Some traits were associated with each other, so the 24 QTL were located on 11 genomic regions. In five QTL located on SSC2, SSC6, and SSC14, each wild boar allele had the effect of increasing types I and IIA muscle fibers and decreasing type IIB muscle fibers. These effects are expected to improve meat quality.     

Quantitative trait loci analysis for growth and carcass traits in a Meishan x Duroc F2 resource population

We constructed a pig F2 resource population by crossing a Meishan sow and a Duroc boar to locate economically important trait loci. The F2 generation was composed of 865 animals (450 males and 415 females) from four F1 males and 24 F1 females and was genotyped for 180 informative microsatellite markers spanning 2,263.6 cM of the whole pig genome. Results of the genome scan showed evidence for significant quantitative trait loci (<1% genomewise error rate) affecting weight at 30 d and average daily gain on Sus scrofa chromosome (SSC) 6, carcass yield on SSC 7, backfat thickness on SSC 7 and SSC X, vertebra number on SSC 1 and SSC 7, loin muscle area on SSC 1 and SSC 7, moisture on SSC 13, intramuscular fat content on SSC 7, and testicular weight on SSC 3 and SSC X. Moreover, 5% genomewise significant QTL were found for birth weight on SSC 7, average daily gain on SSC 4, carcass length on SSC 6, SSC 7, and SSC X and lightness (L value) on SSC 3. We identified 38 QTL for 28 traits at the 5% genomewise level. Of the 38 QTL, 24 QTL for 17 traits were significant at the 1% genomewise level. Analysis of marker genotypes supported the breed of origin results and provided further evidence that a suggestive QTL for circumference of cannon bone also was segregating within the Meishan parent. We identified genomic regions related with growth and meat quality traits. Fine mapping will be required for their application in introgression programs and gene cloning.     

Identification of carcass and meat quality quantitative trait loci in a Landrace pig population selected for growth and leanness

The identification of QTL related to production traits that are relevant for the pig industry has been mostly performed by using divergent crosses. The main objective of the current study was to investigate whether these growth, fatness, and meat quality QTL, previously described in diverse experimental populations, were segregating in a Landrace commercial population selected for litter size, backfat thickness, and growth performance. We have found QTL for carcass weight (posterior P > 0.75), cutlet weight (posterior P > 0.99), weight of ham (posterior P > 0.75), shoulders weight (posterior probability > 0.99), and shear firm-ness (posterior P > 0.99) on pig Chromosome 2. Moreover, QTL with posterior P > 0.75 for fat thickness between the 3rd and 4th ribs (Chromosome 7), rib weights (Chromosome 8), backfat thickness (Chromosomes 8, 9, and 10), and b Minolta color component (Chromosome 7) were identified. These results indicate that commercial purebred populations retain a significant amount of genetic variation, even for traits that have been selected for many generations.     

Bovine quantitative trait loci analysis for growth, carcass, and meat quality traits in an F2 population from a cross between Japanese Black and Limousin

A genome-wide scan for QTL affecting economically important traits in beef production was performed using an F(2) resource family from a Japanese Black x Limousin cross, where 186 F(2) animals were measured for growth, carcass, and meat-quality traits. All family members were genotyped for 313 informative microsatellite markers that spanned 2,382 cM of bovine autosomes. The centromeric region of BTA2 contained significant QTL (i.e., exceeding the genome-wide 5% threshold) for 5 carcass grading traits [LM area, beef marbling standards (BMS) number, luster, quality grade, and firmness), 8 computer image analysis (CIA) traits [LM lean area, ratio of fat area (RFA) to LM area, LM area, RFA to musculus (M.) trapezius area, M. trapezius lean area, M. semispinalis lean area, RFA to M. semispinalis area, and RFA to M. semispinalis capitis area], and 5 meat quality traits (contents of CP, crude fat, moisture, C16:1, and C18:2 of LM). A significant QTL for withers height was detected at 80.3 cM on BTA5. We detected significant QTL for the C14:0 content in backfat and C14:0 and C14:1 content in intermuscular fat around the 62.3 to 71.0 cM region on BTA19 and for C14:0, C14:1, C18:1, and C16:0 content and ratio of total unsaturated fatty acid content to total SFA content in intramuscular fat at 2 different regions on BTA19 (41.1 cM for C14:1 and 62.3 cM for the other 4 traits). Overall, we identified 9 significant QTL regions controlling 27 traits with genome-wide significance of 5%; of these, 22 traits exceeded the 1% genome-wide threshold. Some of the QTL affecting meat quality traits detected in this study might be the same QTL as previously reported. The QTL we identified need to be validated in commercial Japanese Black cattle populations.     

Quantitative trait loci analysis on Sus scrofa chromosome 7 for meat production, meat quality, and carcass traits within a Duroc purebred population

Many QTL analyses related to meat production and meat quality traits have been carried out using an F(2) resource population produced by crossing 2 genetically different breeds. This experiment was intended to investigate whether these QTL were segregating in a purebred Duroc population that had been selected for meat production and meat quality traits during 7 generations. Sus scrofa chromosome 7, for which significant QTL of intramuscular fat and many other traits have already been reported, was studied. The polymorphism of 10 microsatellite markers that were arranged at about 20-cM intervals was investigated on 1,004 pigs. In the selected population, 954 progeny were produced from mating of 99 sires and 286 dams. The QTL analysis for a full-sib family population was examined with the multigeneration pedigree structure of the population. Variance component analysis was used to detect QTL in this population and was examined for the multigeneration pedigree population. In this study, multigenerational pedigree estimated identical by descent coefficients among sibs were produced using Markov chain Monte Carlo methods. The maximum likelihood of odds score was found at the 70-cM position for the LM area, at the 0-cM position for the pork color standard, and at the 120-cM position for the number of thoracic vertebra, but no significant QTL for intramuscular fat were detected on SSC 7. These results indicate that QTL analysis via a variance component method within a purebred population was effective to determine that QTL were segregating in a population of purebred Durocs.     

Economic values for lean meat and fat efficiency in Norwegian Landrace nucleus pig population

A profit function was developed to estimate economic values for new efficiency traits for fattening pigs in Norwegian Landrace. These traits were lean meat efficiency (LME) and fat efficiency (FE). LME and FE described how much feed the animal used to produce 1?kg lean meat and fat as a deviation from the mean. Both traits were derived from analysing total feed consumption in a random regression model which included lean meat and fat content of the carcass (FC) as random covariates. In addition, economic values and breeding values were calculated for days from 40 to 100/120?kg live weight (DAYS), lean meat percentage (LMP) and FC. To compare LME and FE with total feed intake (FI), two indexes were constructed. One index included LME, FE, DAYS, LMP and FC and was referred to as breeding goal A and one index included FI, LMP and DAYS and was referred to as breeding goal B. The standardized economic values for LME and FE were 8.9 and 2.9 EUR/σ_a, respectively. There was a larger variation in the index for breeding goal A (SD?=?5.65) than B (SD?=?3.97). The results suggested that the two efficiency traits had an economic importance in pork production and that there was a potential for increased genetic gain in profit by using breeding goal A compared to breeding goal B.     

Genome-wide mapping and identification of new quantitative trait loci affecting meat production, meat quality, and carcass traits within a Duroc purebred population

Most QTL detection studies in pigs have been carried out in experimental F(2) populations. However, segregation of a QTL must be confirmed within a purebred population for successful implementation of marker-assisted selection. Previously, QTL for meat quality and carcass traits were detected on SSC 7 in a Duroc purebred population. The objectives of the present study were to carry out a whole-genome QTL analysis (except for SSC 7) for meat production, meat quality, and carcass traits and to confirm the presence of segregating QTL in a Duroc purebred population. One thousand and four Duroc pigs were studied from base to seventh generation; the pigs comprised 1 closed population of a complex multigenerational pedigree such that all individuals were related. The pigs were evaluated for 6 growth traits, 7 body size traits, 8 carcass traits, 2 physiological traits, and 11 meat quality traits, and the number of pigs with phenotypes ranged from 421 to 953. A total of 119 markers were genotyped and then used for QTL analysis. We utilized a pedigree-based, multipoint variance components approach to test for linkage between QTL and the phenotypic values using a maximum likelihood method; the logarithm of odds score and QTL genotypic heritability were estimated. A total of 42 QTL with suggestive linkages and 3 QTL with significant linkages for 26 traits were detected. These included selection traits such as daily BW gain, backfat thickness, loin eye muscle area, and intramuscular fat content as well as correlated traits such as body size and meat quality traits. The present study disclosed QTL affecting growth, body size, and carcass, physiological, and meat quality traits in a Duroc purebred population.     

Genome-wide association study for intramuscular fatty acid composition in an Iberian x Landrace cross

The lipid content and fatty acid (FA) profile have an important impact in human health as well as in the technological transformation and nutritional and organoleptic quality of meat. A genome-wide association study (GWAS) on 144 backcross pigs (25% Iberian × 75% Landrace) was performed for 32 traits associated with intramuscular FA composition and indices of FA metabolism. The GWAS was carried out using Qxpak 5.0 and the genotyping information obtained from the Porcine SNP60K BeadChip (Illumina Inc., San Diego, CA). Signals of significant association considering a false- discovery rate (q-value < 0.05) were observed in 15 of the 32 analyzed traits, and a total of 813 trait-associated SNP (TAS), distributed in 43 chromosomal intervals on almost all autosomes, were annotated. According to the clustering analysis based on functional classification, several of the annotated genes are related to FA composition and lipid metabolism. Some interesting positional concordances among TAS and previously reported QTL for FA compositions and/or other lipid traits were also found. These common genomic regions for different traits suggest pleiotropic effects for FA composition and were found primarily on SSC4, SSC8, and SSC16. These results contribute to our understanding of the complex genetic basis of FA composition and FA metabolism.     

Effect of the porcine IGF2-intron3-G3072A substitution in an outbred Large White population and in an Iberian x Landrace cross

The IGF2-intron3-G3072A substitution has been recently described as the causal factor of the imprinted QTL for fat deposition and muscle growth detected within the porcine IGF2 region. The objective of this study was to investigate the IGF2 substitution effect in a Large White outbred population and in an Iberian x Landrace F2 cross. The results showed that the substitution has significant effects on fatness, growth, and shape traits with estimated effects in the expected direction. These results agree with those obtained in the F2 cross, where the IGF2-intron3-G3072A substitution is segregating only in a small family. In addition, a QTL scan has been performed in the F2 population for the traits used in the IGF2 substitution effect validation. Results of this study demonstrated that there are QTL segregating in swine chromosome 2 other than the IGF2 substitution for carcass weight, LM area, and pH measured at 24 h after slaughter. The results confirm the relevance of the IGF2 substitution, but they also show that there are still valuable mutations to be revealed in this chromosome.     

Detection and characterization of quantitative trait loci for meat quality traits in pigs.

In an experimental cross between Meishan and Dutch Large White and Landrace lines, 785 F2 animals with carcass information and their parents were typed for molecular markers covering the entire porcine genome. Linkage was studied between these markers and eight meat quality traits. Quantitative trait locus analyses were performed using interval mapping by regression under two genetic models: 1) the line-cross approach, where the founder lines were assumed to be fixed for different QTL alleles and 2) a half-sib model where a unique allele substitution effect was fitted within each of the 38 half-sib families. The line-cross approach included tests for genomic imprinting and sex-specific QTL effects. In total, three genome-wide significant and 26 suggestive QTL were detected. The significant QTL on chromosomes 3, 4, and 13, affecting meat color, were only detected under the half-sib model. Failure of the line-cross approach to detect the meat color QTL suggests that the founder lines have similar allele frequencies for these QTL. This study provides information on new QTL affecting meat quality traits. It also shows the benefit of analyzing experimental data under different genetic and statistical models.     

Validation of the QTL on SSC4 for meat and carcass quality traits in a commercial crossbred pig population

Porcine chromosome 4 harbours many quantitative trait loci (QTL) affecting meat quality, fatness and carcass composition traits, detected in resource pig populations previously. However, prior to selection in commercial breeds, QTL identified in an intercross between divergent breeds require confirmation, so that they can be segregated. Consequently, the objective of this study was to validate several QTL on porcine chromosome 4 responsible for meat and carcass quality traits. The experimental population consisted of 14 crossbred paternal half-sib families. The region of investigation was the q arm of SSC4 flanked by the markers S0073 and S0813. Regression analysis resulted in the validation of three QTL within the interval: Minolta a * loin, back fat thickness and the weight of trimmed ham. The results were additionally confirmed by factor analysis. Candidate genes were proposed for meat colour, which was the most evident QTL validated in this study.     

Genetic parameter estimation and gene association analyses for meat quality traits in open-air free-range Iberian pigs

Meat quality of Iberian pigs is defined by the combination of their genetic characteristics and the particular production system. To carry out a genetic analysis of the main meat quality traits, we estimated their heritabilities, genetic correlations and the association effects of 32 selected SNPs of 12 candidate genes. A total of ten traits were measured in longissimus dorsi samples from 1,199 Iberian pigs fattened in the traditional free-range system: water holding capacity (thawing, cooking and centrifuge force water losses), instrumental colour (lightness L*, redness a* and yellowness b*), myoglobin content, shear force on cooked meat, and shear force and maximum compression force on dry-cured loin. Estimated heritability values were low to moderate (0.01 to 0.43) being the lowest for L* and the highest for cooking loss. Strong genetic correlations between water holding capacity traits (0.93 to 0.96) and between myoglobin content and a* (0.94) were observed. The association analyses revealed 19 SNPs significantly associated with different traits. Consistent and strong effects were observed between PRKAG3 SNPs (rs319678464G > C and rs330427832C > T), MYH3_rs81437544T > C, CASP3_rs319658214G > T and CTSL_rs332171512A > G and water losses. Also for CAPN1_rs81358667G > A and CASP3_rs319658214G > T and shear force. The SNPs mapping on PRKAG3 showed the highest effects on Minolta colour traits. Genotyping of these SNPs could be useful for the selection of Iberian young boars with similar estimated breeding values for productive traits.     

Quantitative trait loci for behavioural traits in chickens

The detection of quantitative trait loci (QTL) of behavioural traits has mainly been focussed on mouse and rat. With the rapid development of molecular genetics and the statistical tools, QTL mapping for behavioural traits in farm animals is developing. In chicken, a total of 30 QTL involved in pecking-related traits, open-field behaviour, tonic immobility, response to novel objects, and response to a restraint test were detected in different studies. In the search for a useful early predictor for feather pecking (FP) behaviour in adult laying hens, the following was found: FP in young animals is not a predictor for FP in adult animals, however, open-field behaviour in young animals is genetically correlated with FP in adult hens. Before the implementation of FP behaviour or open-field behaviour in breeding programmes, it is essential to know more about the correlation between these behavioural traits and also their relationship with production traits. Nevertheless, with the QTL for behavioural traits and the chicken genome sequence in progress, a better understanding of the underlying genetic mechanisms of behavioural traits will be feasible.     

Meat quality traits were unaffected by a quantitative trait locus affecting leg composition traits in Texel sheep

A QTL affecting leg muscle and fat traits has been identified within the New Zealand Texel population. The QTL maps to a region on OAR 2 with a two-marker haplotype test established at markers BULGE20 and BM81124. These markers encompass the likely position of Growth Differentiation Factor 8 (GDF8). The pleiotropic effects of this QTL on meat quality traits are tested. Objective measures of meat quality including pH, color (L*, a*, and b*), and tenderness (as assessed by Warner-Bratzler shear force measurements) were assessed on longissimus and semi-membranosus muscles of 540 progeny from six Texel sires. Four of these sires were subsequently identified as segregating for leg muscle and fat traits. For these segregating sires, comparison of progeny that had inherited the favorable haplotype from their sire with those that had received the alternate haplotype revealed no significant differences in the meat quality traits assessed. This finding suggests that the muscling QTL does not have pleiotropic effects on meat quality. A general scan for meat quality QTL was carried out using genotype data for eight markers from FCB128 to RM356 flanking 122cM of OAR 2 using Haley-Knott regression. This analysis revealed two QTL for a single sire. A QTL detected in the region of Marker INRA40 for color L* mapped to a site close to the muscling QTL, but there was evidence to suggest it is at a distinct locus. The QTL in the region of Marker RM356 might map distal to Marker RM356, as no peak was observed. This QTL, which seems to affect pH, color a*, color b*, and Warner-Bratzler shear measurements, requires further characterization.     

Ob基因对松辽黑猪肉质和胴体性状的影响

以松辽黑猪为研究对象,分析Ob基因对其肉质和胴体性状的影响。测定30头松辽黑猪的9个肉质性状和6个胴体性状,用PCR-RFLP方法检测Ob基因型。结果显示,松辽黑猪TT基因型平均膘厚比CT基因型低,而肌内脂肪含量、瘦肉率则显著高于后者(P<0.05),其他肉质和胴体性状在Ob基因型间没有显著差异(P>0.05)。结果表明,Ob基因的TT基因型对猪的肉质和胴体性状具有显著的正遗传效应。     

Genetic associations of prolificacy with performance, carcass, meat quality, and leg conformation traits in the Finnish Landrace and Large White pig populations

The objective of this study was to estimate genetic associations of prolificacy traits with other traits under selection in the Finnish Landrace and Large White populations. The prolificacy traits evaluated were total number of piglets born, number of stillborn piglets, piglet mortality during suckling, age at first farrowing, and first farrowing interval. Genetic correlations were estimated with two performance traits (ADG and feed:gain ratio), with two carcass traits (lean percent and fat percent), with four meat quality traits (pH and L* values in longissimus dorsi and semimembranosus muscles), and with two leg conformation traits (overall leg action and buck-kneed forelegs). The data contained prolificacy information on 12,525 and 10,511 sows in the Finnish litter recording scheme and station testing records on 10,372 and 9,838 pigs in Landrace and Large White breeds, respectively. The genetic correlations were estimated by the restricted maximum likelihood method. The most substantial correlations were found between age at first farrowing and lean percent (0.19 in Landrace and 0.27 in Large White), and fat percent (-0.26 in Landrace and -0.18 in Large White), and between number of stillborn piglets and ADG (-0.38 in Landrace and -0.25 in Large White) and feed:gain (0.27 in Landrace and 0.12 in Large White). The correlations are indicative of the benefits of superior growth for piglets already at birth. Similarly, the correlations indicate that age at first farrowing is increasing owing to selection for carcass lean content. There was also clear favorable correlation between performance traits and piglet mortality from birth to weaning in Large White (r(g) was -0.43 between piglet mortality and ADG, and 0.42 between piglet mortality and feed:gain), but not in Landrace (corresponding correlations were 0.26 and -0.22). There was a general tendency that prolificacy traits were favorably correlated with performance traits, and unfavorably with carcass lean and fat percents, whereas there were no clear associations between prolificacy and meat quality or leg conformation. In conclusion, accuracy of estimated breeding values may be improved by accounting for genetic associations between prolificacy, carcass, and performance traits in a multitrait analysis.