Transcriptional analysis of intramuscular fatty acid composition in the longissimus thoracis muscle of Iberian × Landrace back-crossed pigs.

This study aimed at identifying differential gene expression conditional on the fatty acid profile of the longissimus thoracis (Lt) muscle, a prime cut of economic relevance for fresh and cured pork production. A population of 110 Iberian (25%) × Landrace (75%) back-crossed pigs was used, because these two breeds exhibit extreme profiles of intramuscular saturated fatty acid, monounsaturated fatty acid (MUFA) and polyunsaturated fatty acid (PUFA) contents. Total RNA from Lt muscle was individually hybridized to GeneChip Porcine Genome arrays (Affymetrix). A principal component analysis was performed with data from the 110 animals to select 40 extreme animals based on the total fatty acid profile and the MUFA composition (MAP). Comparison of global transcription levels between extreme fatty acid profile pigs (n = 40) resulted in 219 differentially expressed probes (false discovery rate <0.10). Gene ontology, pathway and network analysis indicated that animals with higher percentages of PUFA exhibit a shift toward a more oxidative muscular metabolism state, with a raise in mitochondria function (PPARGC1A, ATF2), fatty acid uptake and oxidation (FABP5, MGLL). On the other hand, 87 probes were differentially expressed between MUFA composition groups (n = 40; false discovery rate <0.10). In particular, muscles rich in n-7 MUFA expressed higher levels of genes involved in lipid metabolism (GLUL, CRAT, PLA2G15) and lower levels of fatty acid elongation genes (ELOVL5). Moreover, the chromosomal position of FABP5, PAQR3, MGLL, PPARGC1A, GLUL and ELOVL5 co-localized with very relevant QTL for fat deposition and composition described in the same resource population. This study represents a complementary approach to identifying genes underlying these QTL effects.

Quantitative trait loci analysis of a Duroc commercial population highlights differences in the genetic determination of meat quality traits at two different muscles.

We performed a whole-genome scan with 110 informative microsatellites in a commercial Duroc population for which growth, fatness, carcass and meat quality phenotypes were available. Importantly, meat quality traits were recorded in two different muscles, that is, gluteus medius (GM) and longissimus thoracis et lumborum (LTL), to find out whether these traits are determined by muscle-specific genetic factors. At the whole-population level, three genome-wide QTL were identified for carcass weight (SSC7, 60 cM), meat redness (SSC13, 84 cM) and yellowness (SSC15, 108 cM). Within-family analyses allowed us to detect genome-wide significant QTL for muscle loin depth between the 3rd and 4th ribs (SSC15, 54 cM), backfat thickness (BFT) in vivo (SSC10, 58 cM), ham weight (SSC9, 69 cM), carcass weight (SSC7, 60 cM; SSC9, 68 cM), BFT on the last rib (SSC11, 48 cM) and GM redness (SSC8, 85 cM; SSC13, 84 cM). Interestingly, there was low positional concordance between meat quality QTL maps obtained for GM and LTL. As a matter of fact, the three genome-wide significant QTL for colour traits (SSC8, SSC13 and SSC15) that we detected in our study were all GM specific. This result suggests that QTL effects might be modulated to a certain extent by genetic and environmental factors linked to muscle function and anatomical location.

Quantitative trait loci for fatness at growing and reproductive stages in Iberian × Meishan F(2) sows.

A considerable number of fatness QTL have been identified in growing pigs, but there is a lack of knowledge about the genetic architecture of this trait in gilts and sows. We have performed a genome scan, in 255 Iberian × Meishan F(2) sows, for backfat thickness (BF) at 150 (BF(150) ) and 210 (BF(210)) days of age, 30 days after conception (BF(30)) and 7-10 days before farrowing (BF(bf)). We have found one BF150 QTL in SSC6 (120 cM) that was highly significant (P < 0.001) at the chromosome-wide level and suggestive at the genome-wide level (P < 0.1). Ten additional chromosome-wide significant QTL were found for sow BF(150) (SSC1, SSC13), BF(210) (SSC6, SSC8, SSC15), BF(30) (SSC5, SSC6) and BF(bf) (SSC1, SSC6, SSC13). The location of several of the BF QTL varied depending on the growing and reproductive status of the sow, suggesting that part of these genetic effects may have a temporal pattern of phenotypic expression.

Expression study on the porcine PTHLH gene and its relationship with sow teat number.

Teat number is an important trait in sows that should accompany the increase in litter size that has been achieved in the last decades through selection. We have previously identified a genome-wide significant QTL for teat number in porcine chromosome SSC5 by means of an experimental Meishan by Iberian F(2) intercross population. In the present report, we have studied the porcine parathyroid hormone-like hormone (PTHLH) gene, which maps to SSC5, as a candidate gene for this trait, as PTHLH is involved in nipple formation during embryogenesis and nipple development during pregnancy and lactation. We have found that porcine PTHLH gene is transcribed into three mRNA species differing in the 5'UTR region. Two of these variants are reported in pigs here for the first time: one was similar to variant 1 described in humans while the other, which was generated by the retention of two small introns, has not been identified before in any other species. In addition, mRNA expression profile for two of the mRNA variants was assessed in 19 pig tissues. Porcine PTHLH showed a widespread expression as it was present in all tested tissues and relative expression of each variant was tissue dependent. Finally, we have performed an association study between a non-synonymous mutation in the coding region of this gene and sow teat number. The PTHLH polymorphism was segregating in our Meishan by Iberian F(2) population at intermediate allelic frequencies. We compared here six different statistical models to choose the one with a better fit and a lower degree of complexity. However, despite the potential negative effect of the PTHLH mutation in the signal peptide of this protein, we did not detect any association between the PTHLH genotype and the sow teat number phenotype, concluding that the causal mutation of the observed QTL is very likely not related to this gene.

Building phenotype networks to improve QTL detection: a comparative analysis of fatty acid and fat traits in pigs.

Models in QTL mapping can be improved by considering all potential variables, i.e. we can use remaining traits other than the trait under study as potential predictors. QTL mapping is often conducted by correcting for a few fixed effects or covariates (e.g. sex, age), although many traits with potential causal relationships between them are recorded. In this work, we evaluate by simulation several procedures to identify optimum models in QTL scans: forward selection, undirected dependency graph and QTL-directed dependency graph (QDG). The latter, QDG, performed better in terms of power and false discovery rate and was applied to fatty acid (FA) composition and fat deposition traits in two pig F2 crosses from China and Spain. Compared with the typical QTL mapping, QDG approach revealed several new QTL. To the contrary, several FA QTL on chromosome 4 (e.g. Palmitic, C16:0; Stearic, C18:0) detected by typical mapping vanished after adjusting for phenotypic covariates in QDG mapping. This suggests that the QTL detected in typical mapping could be indirect. When a QTL is supported by both approaches, there is an increased confidence that the QTL have a primary effect on the corresponding trait. An example is a QTL for C16:1 on chromosome 8. In conclusion, mapping QTL based on causal phenotypic networks can increase power and help to make more biologically sound hypothesis on the genetic architecture of complex traits.

Analysis of candidate genes underlying two epistatic quantitative trait loci on SSC12 affecting litter size in pig.

The previous results from a genome scan for total number of piglets born and number of piglets born alive in a F(2) Iberian by Meishan intercross showed several single and epistatic QTL. One of the most interesting results was obtained for SSC12, where two QTL affecting both traits showed epistatic interaction. In this study, we proposed two genes (SLC9A3R1 and NOS2) as biological and potentially positional candidates underlying these QTL. Both cDNAs were characterized and 23 polymorphisms were detected. A chromosome scan was conducted with 12 markers, plus one SNP in SLC9A3R1 and one in NOS2, covering 110 cM of SSC12. The epistatic QTL (QTL1 at 15 cM and QTL2 at 97 cM) were confirmed, and SLC9A3R1 and NOS2 were mapped around the QTL1 and QTL2 regions respectively. Several SNPs in both genes were tested with standard animal model and marker assisted association tests. The most significant results were obtained with the NOS2 haplotype defined by one missense SNP c.2192C > T (Val to Ala) and a 15 bp duplication at the 3'UTR. This duplication seems to include AU-rich elements, and could be a target site for miRNA, therefore there are statistical and biological indications to consider this haplotype as the potential causal mutation underlying QTL2. SLC9A3R1 results were not conclusive. Although the interaction between the SNPs was not significant, we cannot reject the possibility of interaction of the NOS2 haplotype with other polymorphisms closely linked to the SL9A3R1 SNPs analysed.

A non-synonymous mutation in a conserved site of the MTTP gene is strongly associated with protein activity and fatty acid profile in pigs.

Despite the economic interest of the fatty acid profile in pigs, no gene has been convincingly associated with this trait so far. Here, the porcine microsomal triglyceride transfer protein (MTTP) gene, which plays a crucial role in the assembly of nascent lipoproteins, has been analysed as a positional candidate gene for a QTL affecting the fatty acid composition that was previously identified on chromosome 8 in an Iberian by Landrace F(2) cross. By resequencing a panel of different breeds, a non-synonymous polymorphism in a conserved residue of the lipid transfer domain of MTTP was identified. Association analyses with this polymorphism showed a strong association with the fatty acid composition of porcine fat, much stronger than the QTL effect, in the F(2) cross and in a synthetic Sino-European line. In addition, in vitro activity assays in liver protein extracts have shown that this mutation is also associated with the lipid transfer activity of the MTTP protein (P < 0.1). These results suggest that the detected polymorphism is a potential causal factor of the fatty acid composition QTL. There appears to be an interaction between the porcine MTTP genotype and the type of fat source in the pig diet, which would agree with the previous results on the biology of MTTP biology.

Nucleotide sequence and association analysis of pig apolipoprotein-B and LDL-receptor genes.

Three genes are the major determinants of heritable hypercholesterolemia diseases in humans: APOB, LDLR and LDLRAP1, which encode for proteins that physically interact to promote cholesterol uptake in the cell. We have carried out association analyses of these variants with serum cholesterol and triglycerides concentrations in a half-sib Duroc pig population. Given the structure of the population (six paternal half-sib families), we have used a statistical model that considers separately the allele transmission through dams (at population level) and through sires (within-families from heterozygous sire). Only polymorphisms showing a relevant substitution effect for both male- and female-transmitted alleles are likely to be causal mutations. Thus, although we have found statistical association between genotypes for LDLR and APOB polymorphisms and serum lipid levels (mean allele substitution effects ranging from 15 to 40% of the standard deviation of these traits), none of them seem to be the causal mutation but probably represent closely linked polymorphisms. We have shown here that these three genes also contribute to genetic variability in pigs, with the description of new polymorphisms in their coding regions. Moreover, we have demonstrated that variants on two of these three genes are segregating in a number of commercial breeds. Finally, we report here the coding region for the porcine LDLRAP1 gene and describe a polymorphism in the last exon of this gene.

Genetic parameters and direct, maternal and heterosis effects on litter size in a diallel cross among three commercial varieties of Iberian pig.

The Iberian pig is one of the pig breeds that has the highest meat quality. Traditionally, producers have bred one of the available varieties, exclusively, and have not used crosses between them, which has contrasted sharply with other populations of commercial pigs for which crossbreeding has been a standard procedure. The objective of this study was to perform an experiment under full diallel design among three contemporary commercial varieties of Iberian pig and estimate the additive genetic variation and the crossbreeding effects (direct, maternal and heterosis) for prolificacy. The data set comprised 18 193 records for total number born and number born alive from 3800 sows of three varieties of the Iberian breed (Retinto, Torbiscal and Entrepelado) and their reciprocal crosses (Retinto × Torbiscal, Torbiscal × Retinto, Retinto × Entrepelado, Entrepelado × Retinto, Torbiscal × Entrepelado and Entrepelado × Torbiscal), and a pedigree of 4609 individuals. The analysis was based on a multiple population repeatability model, and we developed a model comparison test that indicated the presence of direct line, maternal and heterosis effects. The results indicated the superiorities of the direct line effect of the Retinto and the maternal effect of the Entrepelado populations. All of the potential crosses produced significant heterosis, and additive genetic variation was higher in the Entrepelado than it was in the other two populations. The recommended cross for the highest yield in prolificacy is a Retinto father and an Entrepelado mother to generate a hybrid commercial sow.

Five genomic regions have a major impact on fat composition in Iberian pigs.

The adipogenic nature of the Iberian pig defines many quality attributes of its fresh meat and dry-cured products. The distinct varieties of Iberian pig exhibit great variability in the genetic parameters for fat deposition and composition in muscle. The aim of this work is to identify common and distinct genomic regions related to fatty acid composition in Retinto, Torbiscal, and Entrepelado Iberian varieties and their reciprocal crosses through a diallelic experiment. In this study, we performed GWAS using a high density SNP array on 382 pigs with the multimarker regression Bayes B method implemented in GenSel. A number of genomic regions showed strong associations with the percentage of saturated and unsaturated fatty acid in intramuscular fat. In particular, five regions with Bayes Factor >100 (SSC2 and SSC7) or >50 (SSC2 and SSC12) explained an important fraction of the genetic variance for miristic, palmitoleic, monounsaturated (>14%), oleic (>10%) and polyunsaturated (>5%) fatty acids. Six genes (RXRB, PSMB8, CHGA, ACACA, PLIN4, PLIN5) located in these regions have been investigated in relation to intramuscular composition variability in Iberian pigs, with two SNPs at the RXRB gene giving the most consistent results on oleic and monounsaturated fatty acid content.

Skew distribution of founder-specific inbreeding depression effects on the longevity of Landrace sows.

Inbreeding is a biological phenomenon of special relevance in domestic species in which its influence has been typically associated with reductions in animal fitness. Nevertheless, recent research suggests substantial founder-specific variability in terms of inbreeding depression on some productive traits, although this centred on a very reduced number of founders. This research focuses on the modelling of founder-specific inbreeding depression (FSID) effects from a more general point of view, characterizing the expected distribution of FSID effects on sow longevity. Under a change-point Weibull proportional hazards model solved through Bayesian inference, a skew-normal a priori distribution for the FSID effects of 19 founders was assumed. In terms of the deviance information criterion, this model was clearly preferred to other prior distributions for FSID effects as well as to a standard analysis of the overall inbreeding depression effect, although all models were consistent with an overall negative genetic effect of inbreeding on sow longevity. The joint posterior distribution of FSID effects showed a skewed pattern with substantial right-tail overexpression, in which the mean (0.036), mode (0.034), S.D. (0.032) and asymmetry parameter (0.045) reported a higher incidence of positive estimates (85.2%) with an unfavourable effect on sow longevity. The founder with the worst inbreeding depression effect reduced sow longevity by 32 days for 1% or 167 days for 10% partial inbreeding. As a whole, our analyses highlighted substantial variability in FSID effects, with unfavourable, neutral and even favourable influences on sow longevity. This heterogeneity could be related to an uneven distribution of the recessive deleterious genetic load among founder genomes, and also with the different selection pressures applied to each founder line. The implementation of skew-normal priors also proved an appealing way to bypass the strict scenario imposed by the standard symmetric-Gaussian distribution, allowing right- and left-tail overexpression as well as non-zero modal estimates.

Bayesian analysis of quantitative traits using skewed distributions.

Statistical models for genetic evaluation often make use of Gaussian distributions. However, some new statistical developments allow the use of an asymmetric distribution for the residuals. Within this context, we analysed three different patterns for the residual term on a data set consisting of 63 208 litter-size records, belonging to 19 255 sows, with a pedigree including 27 911 individuals. The three different residual distributions were: (1) Gaussian distribution, (2) asymmetric Gaussian distribution and (3) asymmetric Gaussian distribution with a hierarchical scheme for the asymmetry parameter. The operational model always included order of parity and herd-year-season as systematic effects, and the permanent environmental and infinitesimal genetic effect of each sow as random effects. The most suitable model using the deviance information criterion (DIC) and posterior predictive checking was model 3. This implies systematic, additive genetic and permanent environmental control of both litter size and the asymmetry parameter of the residual distribution. The asymmetry parameter can be understood as a measure of sensitivity to negative (or positive) environmental influences on phenotypes. The posterior mean (standard deviation) of the additive genetic variance was 0.28 (0.06) for litter size and 0.07 (0.01) for the asymmetry parameter. The posterior mean (standard deviation) of the additive genetic correlation between litter size and the asymmetry parameter was 0.21 (0.07).

An association analysis between a silent C558T polymorphism at the pig vascular cell adhesion molecule 1 locus and sow reproduction and piglet survivability traits.

The vascular cell adhesion molecule 1 (VCAM1) has a strong influence on embryonic development and on the formation of the umbilical cord and placenta. These developmental processes are crucial to ensure the success of pregnancy. In this work, we have identified two T306A and C558T single nucleotide polymorphisms (SNP) at exons 2 and 3 of the pig VCAM1 locus, respectively. The T306A substitution involves a non conservative Asn to Lys replacement at amino acid position 102, whereas the C558T polymorphism is synonymous. An in silico prediction of the consequences of the Asn(102)-->Lys(102) mutation with the PolyPhen software revealed that it is not deleterious. The T306A SNP segregated in the Iberian, Piétrain, Duroc, Large White and Landrace breeds as well as in European wild boars. The C558T SNP also segregated and most of commercial standard breeds. The genotyping of the C558T SNP in an Iberian x Meishan intercross allowed to find a suggestive association (Bonferroni threshold, p < 0.004) between C558T genotype and time the newborn piglet needs to reach the udder (p = 0.013) as well as a significant one with time to make the first ingestion of colostrum (p = 0.003). The biological basis of these associations remains unclear and they should be interpreted with caution.

Genetic parameters and crossbreeding effects of fat deposition and fatty acid profiles in Iberian pig lines.

The aim of this study was to estimate the genetic and environmental parameters and crossbreeding effects on fatty acid and fat traits in the Iberian pig. Our final goal is to explore target selection traits and define crossbreeding strategies. The phenotypes were obtained under intensive management from 470 animals in a diallelic experiment involving Retinto, Torbiscal, and Entrepelado lines. The data set was composed of backfat thickness at the fourth rib (BFT), intramuscular fat (IMF) in the longissimus thoracis (LT), and the fatty acid profile for IMF and subcutaneous fat (SCF) traits. Data were analyzed through a Bayesian bivariate animal model by using a reparameterization of Dickerson's model. The results obtained showed an important genetic determinism for all traits analyzed with heritability ranging from 0.09 to 0.67. The common environment litter effect also had an important effect on IMF (0.34) and its fatty acid composition (0.06-0.53) at slaughter. The additive genetic correlation between BFT and IMF (additive genetic correlation [] = 0.31) suggested that it would be possible to improve lean growth independent of the IMF with an appropriate selection index. Furthermore, the high additive genetic correlation ( = 0.68) found between MUFA tissues would seem to indicate that either the LT or SCF could be used as the reference tissue for MUFA selection. The relevance of the crossbreeding parameters varied according to the traits analyzed. Backfat thickness at the fourth rib and the fatty acid profile of the IMF showed relevant differences between crosses, mostly due to line additive genetic effects associated with the Retinto line. On the contrary, those for IMF crosses were probably mainly attributable to heterosis effects. Particularly, heterosis effects were relevant for the Retinto and Entrepelado crosses (approximately 16% of the trait), which could be valuable for a crossbreeding system involving these lines.

Derivation of a Bayes factor to distinguish between linked or pleiotropic quantitative trait loci.

A simple procedure to calculate the Bayes factor between linked and pleiotropic QTL models is presented. The Bayes factor is calculated from the marginal prior and posterior densities of the locations of the QTL under a linkage and a pleiotropy model. The procedure is computed with a Gibbs sampler, and it can be easily applied to any model including the location of the QTL as a variable. The procedure was compared with a multivariate least-squares method. The proposed procedure showed better results in terms of power of detection of linkage when low information is available. As information increases, the performance of both procedures becomes similar. An example using data provided by an Iberian by Landrace pig intercross is presented. The results showed that three different QTL segregate in SSC6: a pleiotropic QTL affects myristic, palmitic, and eicosadienoic fatty acids; another pleiotropic QTL affects palmitoleic, stearic, and vaccenic fatty acids; and a third QTL affects the percentage of linoleic acid. In the example, the Bayes factor approach was more powerful than the multivariate least-squares approach.

Identification of quantitative trait loci for production traits in commercial pig populations.

The aim of this study was to investigate methods for detecting QTL in outbred commercial pig populations. Several QTL for back fat and growth rate, previously detected in experimental resource populations, were examined for segregation in 10 different populations. Two hundred trait-by-population-by-chromosome tests were performed, resulting in 20 tests being significant at the 5% level. In addition, 53 QTL tests for 11 meat quality traits were declared significant, using a subset of the populations. These results show that a considerable amount of phenotypic variance observed in these populations can be explained by major alleles segregating at several of the loci described. Thus, despite a relatively strong selection pressure for growth and back fat traits in these populations, these alleles have not yet reached fixation. The approaches used here demonstrate that it is possible to verify segregation of QTL in commercial populations by limited genotyping of a selection of informative animals. Such verified QTL may be directly exploited in marker-assisted selection (MAS) programs in commercial populations and their molecular basis may be revealed by positional candidate cloning.

Endometrial gene expression profile of pregnant sows with extreme phenotypes for reproductive efficiency.

Prolificacy can directly impact porcine profitability, but large genetic variation and low heritability have been found regarding litter size among porcine breeds. To identify key differences in gene expression associated to swine reproductive efficiency, we performed a transcriptome analysis of sows' endometrium from an Iberian x Meishan F2 population at day 30-32 of gestation, classified according to their estimated breeding value (EBV) as high (H, EBV > 0) and low (L, EBV < 0) prolificacy phenotypes. For each sample, mRNA and small RNA libraries were RNA-sequenced, identifying 141 genes and 10 miRNAs differentially expressed between H and L groups. We selected four miRNAs based on their role in reproduction, and five genes displaying the highest differences and a positive mapping into known reproductive QTLs for RT-qPCR validation on the whole extreme population. Significant differences were validated for genes: PTGS2 (p = 0.03; H/L ratio = 3.50), PTHLH (p = 0.03; H/L ratio = 3.69), MMP8 (p = 0.01; H/L ratio =4.41) and SCNN1G (p = 0.04; H/L ratio = 3.42). Although selected miRNAs showed similar expression levels between H and L groups, significant correlation was found between the expression level of ssc-miR-133a (p < 0.01) and ssc-miR-92a (p < 0.01) and validated genes. These results provide a better understanding of the genetic architecture of prolificacy-related traits and embryo implantation failure in pigs.

Multivariate analysis of litter size for multiple parities with production traits in pigs: I. Bayesian variance component estimation.

A total of 66,620 records from the first six parities for number of piglets born alive (NBA) from 20,120 Landrace sows and 24,426 records for weight (WT) and backfat thickness (BT) at 175 d of age were analyzed to estimate genetic parameters. The pedigree consisted of 47,186 individuals, including 392 sires and 5,394 dams. Estimates were based on marginal posterior distribution of the genetic parameters obtained using Bayesian inference implemented via the Gibbs sampling procedure with a Data Augmentation step. The posterior means and posterior standard deviation (PSD) for heritability of NBA ranged from 0.064 (PSD 0.005) in the first parity to 0.146 (PSD 0.019) in the sixth parity, always increasing with the order of the parity. The posterior means for genetic correlations of litter size between adjacent parities were, in most cases, greater than 0.80. However, genetic correlation were much lower between nonadjacent parities. For example, the genetic correlation was 0.534 (PSD 0.061) between the fourth and the sixth parity for NBA. The posterior means of heritability for WT and BT were 0.229 (PSD 0.018) and 0.350 (PSD 0.019), respectively. Posterior mean for genetic correlation between WT and BT was 0.339 (PSD 0.044). The posterior means for genetic correlation between production (WT and BT) and reproduction traits (NBA in different parities) were close to zero in most cases. Results from this study suggest that different parities should be considered as different traits. Moreover, selection for growth and backfat should result in no or very little correlated response in litter size.

Multivariate analysis of litter size for multiple parities with production traits in pigs: II. Response to selection for litter size and correlated response to production traits.

Litter size and production trait responses to experimental selection for increased litter size in a Landrace pig population are reported. The numbers of sows and litters available for the first cycle of selection were 3,034 and 961, respectively. Selection was carried out using a BLUP repeatability animal model for number of piglets born alive (NBA). The experiment included one selection and one control line, each with three nonoverlapping generations. The selection line (H) consisted of the 160 sows with the highest breeding values and one boar from each of 25 full-sib families with the highest breeding values. The control line (C) consisted of 160 sows and 25 boars randomly chosen. The two subsequent generations in each line were obtained by random selection. A Bayesian analysis of genetic response using a multivariate model was carried out by Gibbs sampler. Marginal posterior distributions were obtained for direct response in NBA, and for correlated response in weight (WT), and backfat thickness (BT) at 175 d of age. The posterior means and posterior standard deviation (PSD) for direct genetic response of NBA ranged from 0.32 (PSD 0.08) in the first parity to 0.64 (PSD 0.08) in the fourth. The posterior means for correlated genetic response in WT and BT were -0.66 kg (PSD 0.36) and 0.20 mm (PSD 0.10), respectively. For WT and BT, the 95% highest posterior density regions (HPD) contain zero-correlated genetic response. Marginal posterior distributions of selection differentials were investigated. The posterior means for standardized selection differentials for NBA in different parities ranged from 0.70 (PSD 0.12) to 0.94 (PSD 0.06) in females for line H, from 0.22 (PSD 0.19) to 0.34 (PSD 0.10) in males for line H, and from 0.08 (PSD 0.08) to 0.13 (PSD 0.07) in females for line C. All available males were used in line C. Results from this experiment showed that selection for increased litter size is effective. Responses to selection were heterogeneous across parities, suggesting that litter size in each parity may have a different genetic background. No correlated genetic response to growth and backfat thickness was observed.

Correlated response to selection for litter size in pigs: II. Carcass, meat, and fat quality traits.

Data on a pig line selected for litter size (H) and a control line (C) were used to estimate the correlated responses to litter size in carcass, meat, and fat quality traits. The differences between the genetic means of animals from line H and line C were used to estimate correlated responses. No differences were found between the two lines in carcass measurements except backfat depth, which was higher (P < 0.05) in line H (0.69 +/- 0.28 mm). This led to a decrease (P < 0.05) in predicted carcass lean content (-6.0 +/- 2.7 g/kg). Differences in joint weight distribution between lines were primarily due to belly weight, which was higher (P < 0.05) in line H (6.3 +/- 1.2 g/kg). There were no important changes in meat quality traits. Chemical composition of semimembranosus muscle (SM) and subcutaneous backfat (SB) differed between lines only for DM in SB, which was higher (P < 0.05) in line H (15.1 +/- 7.1 mg/g), and for the fatty acid composition of intramuscular fat. The fatty acid profile in line H showed a lower (P < 0.01) proportion of polyunsaturated fatty acids (-14.7 +/- 4.8 mg/g FA), particularly with regard to the content of linoleic acid (-12.5 +/- 3.9 mg/g FA). It is concluded that selection for litter size reduced the lean content in the carcass but the proportion of high-priced cuts and meat quality traits were not affected. However, selection may lead to changes in the composition of intramuscular fat lipids towards a lower content of polyunsaturated fatty acids. The observed correlated effects can be interpreted assuming that selected pigs are more mature at the same weight, though the underlying genetic and physiologic processes that cause them are unknown. The results of this experiment indicate that the metabolic pathways taking part in fat metabolism should be considered first.