Detection of quantitative trait loci for carcass composition traits in pigs

A quantitative trait locus (QTL) analysis of carcass composition data from a three-generation experimental cross between Meishan (MS) and Large White (LW) pig breeds is presented. A total of 488 F2 males issued from six F1 boars and 23 F1 sows, the progeny of six LW boars and six MS sows, were slaughtered at approximately 80 kg live weight and were submitted to a standardised cutting of the carcass. Fifteen traits, i.e. dressing percentage, loin, ham, shoulder, belly, backfat, leaf fat, feet and head weights, two backfat thickness and one muscle depth measurements, ham + loin and back + leaf fat percentages and estimated carcass lean content were analysed. Animals were typed for a total of 137 markers covering the entire porcine genome. Analyses were performed using a line-cross (LC) regression method where founder lines were assumed to be fixed for different QTL alleles and a half/full sib (HFS) maximum likelihood method where allele substitution effects were estimated within each half-/full-sib family. Additional analyses were performed to search for multiple linked QTL and imprinting effects. Significant gene effects were evidenced for both leanness and fatness traits in the telomeric regions of SSC 1q and SSC 2p, on SSC 4, SSC 7 and SSC X. Additional significant QTL were identified for ham weight on SSC 5, for head weight on SSC 1 and SSC 7, for feet weight on SSC 7 and for dressing percentage on SSC X. LW alleles were associated with a higher lean content and a lower fat content of the carcass, except for the fatness trait on SSC 7. Suggestive evidence of linked QTL on SSC 7 and of imprinting effects on SSC 6, SSC 7, SSC 9 and SSC 17 were also obtained.

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title: Genetic mapping of quantitative trait loci for growth and fatness in pigs.
abstract: The European wild boar was crossed with the domesticated Large White pig to genetically dissect phenotypic differences between these populations for growth and fat deposition. The most important effects were clustered on chromosome 4, with a single region accounting for a large part of the breed difference in growth rate, fatness, and length of the small intestine. The study is an advance in genome analyses and documents the usefulness of crosses between divergent outbred populations for the detection and characterization of quantitative trait loci. The genetic mapping of a major locus for fat deposition in the pig could have implications for understanding human obesity.

title: Detection of quantitative trait loci for backfat thickness and intramuscular fat content in pigs (Sus scrofa).
abstract: In an experimental cross between Meishan and Dutch Large White and Landrace lines, 619 F(2) animals and their parents were typed for molecular markers covering the entire porcine genome. Associations were studied between these markers and two fatness traits: intramuscular fat content and backfat thickness. Association analyses were performed using interval mapping by regression under two genetic models: (1) an outbred line-cross model 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 19 half-sib families. Both approaches revealed for backfat thickness a highly significant QTL on chromosome 7 and suggestive evidence for a QTL at chromosome 2. Furthermore, suggestive QTL affecting backfat thickness were detected on chromosomes 1 and 6 under the line-cross model. For intramuscular fat content the line-cross approach showed suggestive evidence for QTL on chromosomes 2, 4, and 6, whereas the half-sib analysis showed suggestive linkage for chromosomes 4 and 7. The nature of the QTL effects and assumptions underlying both models could explain discrepancies between the findings under the two models. It is concluded that both approaches can complement each other in the analysis of data from outbred line crosses.

title: A QTL for intramuscular fat and backfat thickness is located on porcine Chromosome 6
abstract: populations coming from crosses of divergentbreeds. Many of these experiments have succeeded in identifyingchromosomal regions associated with growth, backfat, and repro-duction (see Rothschild and Plastow 1999 for a review). Nonethe-less, and despite its increasing economic importance and the num-ber of ongoing QTL experiments, the information concerning meatquality traits is much scarcer.We have developed an F

title: Identification of quantitative trait loci affecting carcass composition in swine: I. Fat deposition traits.
abstract: One of the major determining factors in the price of market hogs today is backfat depth. Therefore, identification of regions of the genome affecting this trait could be very useful to the swine industry. A large resource population (n = 540) was developed by backcrossing F1 Meishan-White composite females to either Meishan or White composite boars. A genomic scan was conducted by genotyping all animals with microsatellite markers spaced at approximately 20-cM intervals across the entire porcine genome. Breed of origin for all chromosomal segments was determined using multipoint linkage procedures, and a least squares regression analysis was conducted. Nominal P-values were converted to a genome-wide level of significance to adjust for the numerous tests conducted. Traits analyzed were backfat depths at the first rib (FRIB); 10th rib (10th); last rib (LRIB); last lumbar (LLUM) vertebra; the average of FRIB, LRIB, and LLUM (AVBF); and weight of leaf fat (LEAF). Significant quantitative trait loci (QTL) were detected on chromosomes 1, 7, and X. Suggestive evidence for QTL was present on chromosomes 5, 8, 9, 10, 13, and 14. Genotypic means for the loci detected indicated a predominantly additive mode of inheritance. Meishan alleles produced fatter pigs for all loci except those on chromosomes 7 and 10. Additional research should be conducted to refine the estimated position of each QTL and its effect and determine epistatic interactions. These loci should be evaluated in other germplasms to determine whether allelic variation at the QTL exists in other breeds.

title: Identification of quantitative trait loci affecting carcass composition in swine: II. Muscling and wholesale product yield traits.
abstract: A genomic scan was conducted on 540 reciprocal backcross Meishan x White composite pigs for hot carcass weight (HCWT); loin eye area (LOIN); carcass length (CRCL); belly weight (BELLY); and weight of trimmed ham, loin, picnic, and Boston butt adjusted to a constant live (TWPLWT) or carcass (TWPCWT) weight. Genetic markers spanned the entire porcine linkage map and were spaced at approximately 20-cM intervals. Grandparental breed of origin for all chromosomal segments was determined using multipoint linkage procedures, and a least squares regression analysis was conducted. Nominal P-values were converted to a genome-wide level of significance to adjust for the number of tests actually conducted. Seven associations were significant at the genome-wide level relating to chromosomes 1 (SSC 1), 7 (SSC 7), and X (SSC X). The SSC 1 region affected LOIN, TWPLWT, and TWPCWT; SSC 7 affected HWCT and CRCL; and SSC X affected TWPLWT and TWPCWT. Twelve associations relating to seven chromosomal regions (including SSC 1 and X) presented suggestive evidence for quantitative trait loci (QTL), and many of these regions are likely to contain QTL. Chromosomes 8 and 14 had two and three traits with suggestive evidence for QTL, respectively. Many pleiotropic effects were detected for regions on SSC 1, 7, 14, and X in this study and a companion study looking for fat deposition QTL in the same population. In addition, SSC 4 was nearly significant for CRCL in the same region identified as affecting backfat in a wild boar x Large White population. These results expand our knowledge of the inheritance of quantitative traits and are directly relevant to composite populations containing Meishan germplasm.

title: Mapping of quantitative trait loci on porcine chromosome 4.
abstract: A F2 population derived from a cross between European Large White and Chinese Meishan pigs was established in order to study the genetic basis of breed differences for growth and fat traits. Chromosome 4 was chosen for initial study as previous work had revealed quantitative trait loci (QTLs) on this chromosome affected growth and fat traits in a Wild Boar x Large White cross. Individuals in the F2 population were typed for nine markers spanning a region of approximately 124 CM. We found evidence for QTLs affecting growth between weaning and the end of test (additive effect: 43.4 g/day) and fat depth measured in the mid-back position (additive effect: 1.82 mm). There was no evidence of interactions between the QTLs and sex, grandparents or F1 sires, suggesting that the detected QTLs were fixed for alternative alleles in the Meishan and Large White breeds. Comparison of locations suggests that these QTLs could be the same as those found in the Wild Boar x Large White cross.

title: Detection of quantitative trait loci for growth and fatness in pigs
abstract: A quantitative trait locus (QTL) analysis of growth and fatness data from a three-generation experimental cross between Meishan (MS) and Large White (LW) pig breeds is presented. Six boars and 23 F1 sows, the progeny of six LW boars and six MS sows, produced 530 F2 males and 573 F2 females. Nine growth traits, i.e. body weight at birth and at 3, 10, 13, 17 and 22 weeks of age, average daily gain from birth to 3 weeks, from 3 to 10 weeks and from 10 to 22 weeks of age, as well as backfat thickness at 13, 17 and 22 weeks of age and at 40 and 60 kg live weight were analysed. Animals were typed for a total of 137 markers covering the entire porcine genome. Analyses were performed using two interval mapping methods: a line-cross (LC) regression method where founder lines were assumed to be fixed for different QTL alleles and a half-/full-sib (HFS) maximum likelihood method where allele substitution effects were estimated within each half-/full-sib family. Both methods revealed highly significant gene effects for growth on chromosomes 1, 4 and 7 and for backfat thickness on chromosomes 1, 4, 5, 7 and X, and significant gene effects on chromosome 6 for growth and backfat thickness. Suggestive QTLs were also revealed by both methods on chromosomes 2 and 3 for growth and 2 for backfat thickness. Significant gene effects were detected for growth on chromosomes 11, 13, 14, 16 and 18 and for backfat thickness on chromosome 8, 10, 13 and 14. LW alleles were associated with high growth rate and low backfat thickness, except for those of chromosome 7 and to a lesser extent early-growth alleles on chromosomes 1 and 2 and backfat thickness alleles on chromosome 6.

title: Recovery of inter-block information when block sizes are unequal
abstract: A method is proposed for estimating intra-block and inter-block weights in the analysis of incomplete block designs with block sizes not necessarily equal. The method consists of maximizing the likelihood, not of all the data, but of a set of selected error contrasts. When block sizes are equal results are identical with those obtained by the method of Nelder (1968) for generally balanced designs. Although mainly concerned with incomplete block designs the paper also gives in outline an extension of the modified maximum likelihood procedure to designs with a more complicated block structure. In this paper we consider the estimation of weights to be used in the recovery of interblock information in incomplete block designs with possibly unequal block sizes. The problem can also be thought of as one of estimating constants and components of variance from data arranged in a general two-way classification when the effects of one classification are regarded as fixed and the effects of the second classification are regarded as random. Nelder (1968) described the efficient estimation of weights in generally balanced designs, in which the blocks are usually, although not always, of equal size. Lack of balance resulting from unequal block sizes is, however, common in some experimental work, for example in animal breeding experiments. The maximum likelihood procedure described by Hartley & Rao (1967) can be used but does not give the same estimates as Nelder's method in the balanced case. As will be shown, the two methods in effect use the same weighted sums of squares of residuals but assign different expectations. In the maximum likelihood approach, expectations are taken over a conditional distribution with the treatment effects fixed at their estimated values. In contrast Nelder uses unconditional expectations. The difference between the two methods is analogous to the well-known difference between two methods of estimating the variance o2 of a normal distribution, given a random sample of n values. Both methods use the same total sum of squares of deviations.

title: Mapping quantitative trait loci in crosses between outbred lines using least squares.
abstract: The use of genetic maps based upon molecular markers has allowed the dissection of some of the factors underlying quantitative variation in crosses between inbred lines. For many species crossing inbred lines is not a practical proposition, although crosses between genetically very different outbred lines are possible. Here we develop a least squares method for the analysis of crosses between outbred lines which simultaneously uses information from multiple linked markers. The method is suitable for crosses where the lines may be segregating at marker loci but can be assumed to be fixed for alternative alleles at the major quantitative trait loci (QTLs) affecting the traits under analysis (e.g., crosses between divergent selection lines or breeds with different selection histories). The simultaneous use of multiple markers from a linkage group increases the sensitivity of the test statistic, and thus the power for the detection of QTLs, compared to the use of single markers or markers flanking an interval. The gain is greater for more closely spaced markers and for markers of lower information content. Use of multiple markers can also remove the bias in the estimated position and effect of a QTL which may result when different markers in a linkage group vary in their heterozygosity in the F1 (and thus in their information content) and are considered only singly or a pair at a time. The method is relatively simple to apply so that more complex models can be fitted than is currently possible by maximum likelihood. Thus fixed effects of background genotype can be fitted simultaneously with the exploration of a single linkage group which will increase the power to detect QTLs by reducing the residual variance. More complex models with several QTLs in the same linkage group and two-locus interactions between QTLs can similarly be examined. Thus least squares provides a powerful tool to extend the range of crosses from which QTLs can be dissected whilst at the same time allowing flexible and realistic models to be explored.

title: Multiple marker mapping of quantitative trait loci in a cross between outbred wild boar and large white pigs.
abstract: A quantitative trait locus (QTL) analysis of growth and fatness data from a three generation pig experiment is presented. The population of 199 F2 animals was derived from a cross between wild boar and Large White pigs. Animals were typed for 240 markers spanning 23 Morgans of 18 autosomes and the X chromosome. A series of analyses are presented within a least squares framework. First, these identify chromosomes containing loci controlling trait variation and subsequently attempt to map QTLs to locations within chromosomes. This population gives evidence for a large QTL affecting back fat and another for abdominal fat segregating on chromosome 4. The best locations for these QTLs are within 4 cM of each other and, hence, this is likely to be a single QTL affecting both traits. The allele inherited from the wild boar causes an increase in fat deposition. A QTL for intestinal length was also located in the same region on chromosome 4 and could be the same QTL with pleiotropic effects. Significant effects, owing to multiple QTLs, for intestinal length were identified on chromosomes 3 and 5. A single QTL affecting growth rate to 30 kg was located on chromosome 13 such that the Large White allele increased early growth rate, another QTL on chromosome 10 affected growth rate from 30 to 70 kg and another on chromosome 4 affected growth rate to 70 kg.

title: Genetic dissection of complex traits: guidelines for interpreting and reporting linkage results
abstract: Genetic studies are under way for many complex traits, spurred by the recent feasibility of whole genome scans. Clear guidelines for the interpretation of linkage results are needed to avoid a flood of false positive claims. At the same time, an overly cautious approach runs the risk of causing true hints of linkage to be missed. We address this problem by proposing specific standards designed to maintain rigor while also promoting communication.

title: Empirical threshold values for quantitative trait mapping.
abstract: The detection of genes that control quantitative characters is a problem of great interest to the genetic mapping community. Methods for locating these quantitative trait loci (QTL) relative to maps of genetic markers are now widely used. This paper addresses an issue common to all QTL mapping methods, that of determining an appropriate threshold value for declaring significant QTL effects. An empirical method is described, based on the concept of a permutation test, for estimating threshold values that are tailored to the experimental data at hand. The method is demonstrated using two real data sets derived from F(2) and recombinant inbred plant populations. An example using simulated data from a backcross design illustrates the effect of marker density on threshold values.

title: Growth, carcass and meat quality performance of crossbred pigs with graded proportions of Meishan genes
abstract: Growth, carcass and meat quality traits were measured in 2 different experimental herds on male and female pigs produced from matings between Pietrain boars and 12 genetic types of sows with graded proportions of Large White (LW) and Meishan (MS) genes. Growth records (from 30-100 kg liveweight) were obtained on ad libitum feeding on a total of 1 640 pigs, among which 1 200 were submitted to carcass evaluation and meat quality measurements. Genetic type mean performance essentially varied according to the relative proportions of MS and LW genes in the dam and could hence be characterized by a single parameter, difference in crossbreeding (!;yls_LW)! which measures the difference between MS and LW breeds used as dam breeds. Differences in crossbreeding were unfavourable to MS for all growth and carcass traits. Average estimates of .&eth;. MSLWwere -71±16 g/d; 0.21!0.07; -2.4!0.3% ; -9.0±0.5% for average daily gain (ADG) feed conversion ratio, killing out percentage and estimated carcass lean content (% M), respectively. However, significant herd differences were observed for ADG and %M. The 2 herd estimates were -51± 16 g/d and -92::!::30 g/d for ADG, -7.3±0.6% and 10. 7 + 1.5%, for %M. Conversely, differences in crossbreeding for meat quality traits were in favour of MS, with an advantage of 1.1 t 0.4 point in meat quality index over LW, ie one third of a phenotypic standard deviation.

title: Identification of quantitative trait loci affecting birth characters and accumulation of backfat and weight in a Meishan-White Composite resource population.
abstract: A search for genomic regions affecting birth characters and accretion of weight and backfat was conducted in a Meishan-White Composite reciprocal backcross resource population. Birth traits analyzed (n = 750) were vigor score, number of nipples, and birth weight. Subsequent measures on gilts and barrows (n = 706) analyzed were weaning weight, 8-wk weight, ADG from 8 to 18 wk of age, ADG from 18 to 26 wk of age, 26-wk weight, and backfat over the first rib, last rib, and last lumbar vertebrae at 14 and 26 (n = 599) wk of age. Feed intake and growth of 92 individually penned barrows were also analyzed. A genomic scan was conducted with microsatellite markers spaced at approximately 20-cM intervals, a least squares regression interval analysis was implemented, and significance values were converted to genomewide levels. No associations were detected for traits measured at birth except for number of nipples, where one significant and two suggestive regions were identified on chromosomes (SSC) 10, 1, and 3, respectively. Early growth was affected by a region on SSC 1 as evidenced by associations with weights collected at weaning and 8 wk of age and ADG from 8 to 18 wk of age. Other regions detected for early growth rate were on SSC 2, 12, and X. Chromosomal regions on SSC 6 and 7 affected ADG from 18 to 26 wk of age. All measures of backfat were affected by regions on SSC 1 and X, whereas SSC 7 consistently affected backfat measures recorded at 26 wk of age. Suggestive evidence for QTL affecting backfat at 14 wk of age was also detected on SSC 2, 6, 8, and 9. These results have improved our knowledge about the genetics of growth rate and fat accretion at the molecular level in swine.

title: Interval mapping of growth in divergent swine cross
abstract: A genomic scan of 18 swine autosomal chromosomes was constructed with 119 polymorphic microsatellite (ms) markers to identify quantitative trait loci (QTL) for 11 growth traits in the University of Illinois Meishan × Yorkshire Swine Resource Family. A significant QTL effect was found for post-weaning average daily gain (ADG) between 5.5 and 56 kg of body weight that mapped between markers SW373 and SW1301 near the telomere of Chromosome (Chr) 1 q (SSC1). This QTL effect had a nominal (pointwise) p-value of 0.000007, a genome wide p-value of 0.012, and accounted for 26% of the F2 phenotypic variance. The same chromosome region also had significant effects on ADG between birth and 56 kg body weight (p-value = .000227), and on ADG between 35 and 56 kg (p-value = .00077). These observations suggest that a significant QTL for post-weaning growth resides on SSC1.

title: Quantitative trait loci (QTL) analysis in a Meishan x Göttingen cross population.
abstract: In order to locate the genetic regions in the swine genome that are responsible for economically important traits, a resource population has been constructed by mating two female Meishan pigs with a male Göttingen miniature pig. In subsequent generations, 265 F2 offspring were produced from two F1 males and 19 F1 females. The F2 offspring were scored for eight traits including growth rate, teat number, vertebra number and backfat thickness, and genotyped for 318 genetic markers spanning the swine genome. Least-square analysis revealed quantitative trait loci (QTL) effects for vertebra number on chromosomes 1 and 2; for teat number on chromosomes 1 and 7; for birth weight on chromosome 1; for average daily gain between 4 and 13 weeks of age on chromosomes 9 and 10; for backfat thickness on chromosome 7; and for backskin thickness on chromosome 3.

title: Identification of porcine Lhx3 and SF1 as candidate genes for QTL affecting growth and reproduction traits in swine.
abstract: The distal portion of the long arm of porcine chromosome 1 has been shown to harbour several quantitative trait loci affecting growth and reproductive traits in swine. In order to identify potential candidate genes that might underlie these effects, a comparative mapping analysis was undertaken to define the extent of orthologous segments of human chromosome 9. A microsatellite associated with heat shock protein (HSP) A5 was used to define the proximal boundary of the quantitative trait loci (QTL) region, which suggests the human orthologue of the gene(s) responsible for the observed effects lies between HSPA5 and the q arm telomere of human chromosome 9. Examination of this region revealed two candidate genes with known roles in production of hormones essential to growth and reproductive function. The steroidogenic factor 1 and Lhx3 LIM homeodomain transcription factor genes were mapped to 123 and 155 cM, respectively, of the Sus scrofa chromosome 1 (SSC1) linkage group, placing both genes within the confidence interval for the observed QTL. To further evaluate Lhx3, we examined the expression profile during porcine embryonic development. Low levels were detected at early embryonic stages, when development of the nervous system is proceeding. A transient increase in expression level is observed during the time of pituitary organogenesis and again at the time of differentiation of anterior pituitary cells, with relatively high levels of expression persisting in the adult pituitary gland. This ontology is consistent with Lhx3 being a candidate gene for the QTL.

title: A paternally expressed QTL affecting skeletal and cardiac muscle mass in pigs maps to the IGF2 locus
abstract: A paternally expressed QTL affecting skeletal and cardiac muscle mass in pigs maps to the IGF2 locus

title: An imprinted QTL with major effect on muscle mass and fat deposition maps to the IGF2 locus in pigs
abstract: An imprinted QTL with major effect on muscle mass and fat deposition maps to the IGF2 locus in pigs

title: Mapping Quantitative Trait Loci for Carcass and Meat Quality Traits in a Wild Boar × Large White Intercross 1
abstract: An intercross between wild boar and a domestic Large White pig population was used to map quantitative trait loci (QTL) for body proportions, weight of internal organs, carcass composition, and meat quality. The results concerning growth traits and fat deposition traits have been reported elsewhere. In the present study, all 200 F2 animals, their parents, and their grandparents were genotyped for 236 markers. The marker genotypes were used to calculate the additive and dominance coefficients at fixed positions in the genome of each F2 animal, and the trait values were regressed onto these coefficients in intervals of 1 cM. In addition, the effect of proportion of wild boar alleles was tested for each chromosome. Significant QTL effects were found for percentage lean meat and percentage lean meat plus bone in various cuts, proportion of bone in relation to lean meat in ham, muscle area, and carcass length. The significant QTL were located on chromosomes 2, 3, 4, and 8. Each QTL explained 9 to 16% of the residual variance of the traits. Gene action for most QTL was largely additive. For meat quality traits, there were no QTL that reached the significance threshold. However, the average proportion of wild boar alleles across the genome had highly significant effects on reflectance and drip loss. The results show that there are several chromosome regions with a considerable effect on carcass traits in pigs.

title: Fine mapping and imprinting analysis for fatness trait QTLs in pigs
abstract: Quantitative trait loci (QTL) for fatness traits were reported recently in an experimental Meishan × Large White and Landrace F2 cross. To further investigate the regions on pig Chr 2 (SSC2), SSC4, and SSC7, 25 additional markers from these regions were typed on 800 animals (619 F2 animals, their F1 parents, and F0 grandfathers). Compared with the published maps, a modified order of markers was observed for SSC4 and SSC7. QTL analyses were performed both within the half-sib families as well as across families (line cross). Furthermore, a QTL model accounting for imprinting effects was tested. Information content could be increased considerably on all three chromosomes. Evidence for the backfat thickness QTL on SSC7 was increased, and the location could be reduced to a 33-cM confidence interval. The QTL for intramuscular fat on SSC4 could not be detected in this half-sib analysis, whereas under the line cross model a suggestive QTL on a different position on SSC4 was detected. For SSC2, in the half-sib analysis, a suggestive QTL for backfat thickness was detected with the best position at 26 cM. Imprinting analysis, however, revealed a genome-wise, significant, paternally expressed QTL on SSC2 with the best position at 63 cM. Our results suggest that this QTL is different from the previously reported paternally expressed QTL for muscle mass and fat deposition on the distal tip of SSC2p.

title: A directed search for quantitative trait loci on chromosomes 4 and 7 in pigs.
abstract: Improvements in the porcine genetic map and availability of resource families to study performance traits in pigs have made it possible to re-examine previous findings that linked certain traits to genes or chromosomal regions. Previous studies suggested that chromosomes 4 and 7 may be associated with growth and performance traits. To confirm these previous results, an interval mapping-regression approach was used to determine whether quantitative trait loci (QTL) exist in the Iowa State University reference/resource families. Traits measured were birth weight; body weight at 21 d; weaning weight (weight at 42 d); average daily gain; backfat at the first, 10th, lumbar, and last ribs and average backfat thickness; loin eye area; meat color; marbling; and firmness. The total number of F2 pigs used ranged from 241 to 330 and came from five Chinese x American resource families. Five markers (S0001, SW871, S0175, S0214, and SW445) were genotyped and mapped on chromosome 4, and so were 10 markers (S0064, tumor necrosis factor alpha [TNFalpha], S0102, S0078, S0158, S0066, SW304, SW1083, S0101, and S0212) on chromosome 7. Data were analyzed for each family (breed cross) separately and were also pooled. Experiment-wise thresholds were used to determine significance. Suggestive evidence of QTL on chromosomes 4 and 7 was observed for several traits in pooled and individual family analyses. Suggestive evidence of a QTL with a relatively large effect for average daily gain was detected on chromosome 4 in the pooled analysis. Significant (P < .05) evidence for QTL was seen on chromosome 7 for 10th-rib, last-rib, and average backfat thickness in the pooled data set in a region of the chromosome that was near TNFalpha. These results verify in part that chromosomes 4 and 7 contain QTL for growth and carcass traits.

title: The adipocyte fatty acid-binding protein locus: characterization and association with intramuscular fat content in pigs
abstract: The porcine A-FABP gene (FABP4) was isolated and sequenced to study the role of A-FABP in the differentiation of intramuscular fat (IMF) accretion in pigs. The coding sequence of the porcine A-FABP gene is highly conserved across human, mouse, and rat. Moreover, all the functionally important amino acids are conserved. This high similarity extends into the first 270 bp of the 5′ upstream region. Within this region, a 56-bp nucleotide sequence was completely identical with the corresponding sequence in the mouse A-FABP gene, which contains the transcription factor binding sites for C/EBP and AP-1, and is implicated in the differentiation-dependent regulation of A-FABP. The A-FABP gene was assigned to porcine Chromosome (Chr) 4 by a porcine sequence-specific PCR on a cell hybrid panel, fully consistent with comparative mapping data with human and mouse. In the first intron of the porcine A-FABP gene, a microsatellite sequence was detected that was polymorphic for all six pig breeds tested. This genetic variation within the A-FABP gene was associated with differences in IMF content and possibly growth in a Duroc population, whereas no effect on backfat thickness and drip loss of the meat were detected. A considerable and significant contrast of approximately 1% IMF was observed between certain genotype classes. We conclude that the A-FABP locus is involved in the regulation of intramuscular fat accretion in Duroc pigs.

title: The effect of adipocyte and heart fatty acid-binding protein genes on intramuscular fat and backfat content in Meishan crossbred pigs.
abstract: Effects of genetic variation in porcine adipocyte and heart fatty acid-binding protein genes, A-FABP and H-FABP, respectively, on intramuscular fat (IMF) content and backfat thickness (BFT) were examined in F2 crossbreds of Meishan and Western pigs. The involvement of each FABP gene in IMF accretion was studied to confirm previous results for Duroc pigs. The F2 crossbred pigs were genotyped for various markers including microsatellite sequences situated within both FABP genes. Linkage analysis assigned the A-FABP and H-FABP genes to marker intervals S0001-S0217 (20 cM) on SSC4 and Sw316-S0003 (16.6 cM) on SSC6, respectively, refining previous chromosomal assignments. Next, the role of both chromosome regions/genes on genetic variation in IMF content and BFT was studied by 1) screening SSC4 and SSC6 for QTL affecting both traits by performing a line-cross analysis and 2) estimation of the effect of individual A-FABP and H-FABP alleles on both traits. In the first analysis, suggestive and chromosome-wise significant evidence for a QTL affecting IMF was detected on SSC6. The H-FABP gene is a candidate gene for this effect because it resides within the large region containing this putative QTL. The second analysis showed a considerable but nonsignificant effect of H-FABP microsatellite alleles on IMF content. Suggestive evidence for a QTL affecting BFT was found on SSC6, but H-FABP was excluded as a candidate gene. In conclusion, present and previous results support involvement of H-FABP gene polymorphisms in IMF accretion independently from BFT in pigs. Therefore, implementation of these polymorphisms in marker-assisted selection to control IMF content independently from BFT may be considered. In contrast to previous findings for Duroc pigs, no evidence was found for an effect of the A-FABP gene on IMF or BFT in this population.

title: The X Chromosome harbors quantitative trait loci for backfat thickness and intramuscular fat content in pigs
abstract: Genetic markers have been used in experimental crosses in pigs todissect genetic variation in quantitative traits (e.g., Andersson et al.1994; Knott et al. 1998; Rohrer and Keele 1998). We reportedrecently on the search for autosomal quantitative trait loci (QTL)for fatness traits in an experimental intercross between the obeseChinese Meishan breed and lean Dutch White production lines (DeKoning et al. 1999; Rattink et al. 2000). In this paper, the analysisto look for QTL on the X Chromosome (Chr) in this cross ispresented.Briefly, 19 Meishan boars were mated to 120 sows of Whitelines from five Dutch breeding companies. From the F

title: A molecular genome scan analysis to identify chromosomal regions influencing economic traits in the pig. I. Growth and body composition
abstract: Genome scans can be employed to identify chromosomal regions and eventually genes (quantitative trait loci or QTL) that control quantitative traits of economic importance. A three-generation resource family was developed by using two Berkshire grand sires and nine Yorkshire grand dams to detect QTL for growth and body composition traits in pigs. A total of 525 F2 progeny were produced from 65 matings. All F2 animals were phenotyped for birth weight, 16-day weight, growth rate, carcass weight, carcass length, back fat thickness, and loin eye area. Animals were genotyped for 125 microsatellite markers covering the genome. Least squares regression interval mapping was used for QTL detection. All carcass traits were adjusted for live weight at slaughter. A total of 16 significant QTL, as determined by a permutation test, were detected at the 5% chromosome-wise level for growth traits on Chromosomes (Chrs) 1, 2, 3, 4, 6, 7, 8, 9, 11, 13, 14, and X, of which two were significant at the 5% genome-wise level and two at the 1% genome-wise level (on Chrs 1, 2, and 4). For composition traits, 20 QTL were significant at the 5% chromosome-wise level (on Chrs 1, 4, 5, 6, 7, 12, 13, 14, 18), of which one was significant at the 5% genome-wise level and three were significant at the 1% genome-wise level (on Chrs 1, 5, and 7). For several QTL the favorable allele originated from the breed with the lower trait mean.

title: Mapping quantitative trait loci for principal components of bone measurements and osteochondrosis scores in a wild boar x large white intercross.
abstract: Data on osteochondrosis and femur dimensions from 195 F2 pigs from a wild boar x Large White intercross were analysed with the aim of detecting quantitative trait loci (QTLs) for normal and disturbed bone formation. The information from numerous recorded traits was summarized by principal component analysis and analysed by least-squares interval mapping. An increase in the proportion of wild boar alleles across the genome increased length versus width of femur and reduced the prevalence of osteochondrosis. The presence of QTLs with an impact on femur dimensions was indicated on chromosomes 2, 4, 16 and 17 and on osteochondrosis on chromosomes 5, 13 and 15. A substantial effect of the chromosome 5 QTL calls for further studies within commercial populations to evaluate whether marker-assisted selection could be used to reduce the prevalence of osteochondrosis.

title: Effects of exogenous porcine somatotropin (pST) administration on growth performance, carcass traits, and pork meat quality of Meishan, Pietrain, and crossbred gilts.
abstract: Seventeen to twenty-three females of lean (Pietrain, PI), fat (Meishan, MS), or intermediate genotype (PI x [3/4 Large White x 1/4 MS]), referred to as crossbred (CR), were injected between 60 and 100 kg live weight with 6 mg/d of porcine somatotropin (pST) and compared to similar numbers of control females receiving the vehicle only. Average daily gain increased similarly in the three genotypes (125 g/d). Feed conversion ratio tended to decrease to a higher extent in MS (-2.0 kg of feed/kg of gain) than in the other two genotypes (-1.1 and -.9 kg of feed/kg of gain for CR and PI, respectively). A significant genotype x treatment interaction was also observed for backfat thickness (BF) and fat, muscle, and bone development. Effects of pST in PI, CR, and MS pigs were, respectively, -6.2, -9.6, and -16.1 mm for BF and 3.0, 6.8, and 11.8% carcass muscle. The influence of pST on physical measurements of meat quality was rather low, although desirable effects (P less than .05) were obtained on the reflectance and water-holding capacity of PI and CR. Intramuscular fat content was reduced by approximately 1% in MS and CR but not in PI. The metatarsals of pST-treated animals had a higher water content (except in PI), a lower mineralization, and a lower breaking strength (except in MS). The existence of breed variations in the response to pST might result in changes of the relative merit of crossbreeding schemes.

title: A further look at quantitative trait loci affecting growth and fatness in a cross between Meishan and Large White pig populations
abstract: A detailed quantitative trait locus (QTL) analysis of growth and fatness data from a three generation experimental cross between Large White (LW) and Meishan (MS) pig breeds was carried out to search for sex × QTL interactions, imprinting effects and multiple linked QTLs. A total of 530 F2 males and 573 F2 females issued from 6 F1 boars and 23 F1 sows were typed for a total of 137 markers covering the entire porcine genome. Nine growth traits and three backfat thickness measurements were analysed. All analyses were performed using line cross regression procedures. A QTL with sex-specific expression was revealed in the proximal region of chromosome 8, although some confusion between herd and sex effects could not be discarded. This previously undetected QTL affected male growth during the fattening period, with a favourable additive effect of the LW allele. The analyses also revealed the presence of two linked QTLs segregating on chromosome 1, affecting growth traits during the post-weaning period. The first QTL, previously detected using a single QTL model, was located at the end of the q arm of chromosome 1 and had a favourable MS allele. The second QTL had a favourable LW allele and was located in the proximal extremity of the q arm of chromosome 1. Suggestive genomic imprinting was found in the distal region of chromosome 9 affecting growth during the fattening period.

title: Genome-wide scan for body composition in pigs reveals important role of imprinting.
abstract: The role of imprinting in body composition was investigated in an experimental cross between Chinese Meishan pigs and commercial Dutch pigs. A whole-genome scan revealed significant evidence for five quantitative trait loci (QTL) affecting body composition, of which four were imprinted. Imprinting was tested with a statistical model that separated the expression of paternally and maternally inherited alleles. For back fat thickness, a paternally expressed QTL was found on Sus scrofa chromosome 2 (SSC2), and a Mendelian-expressed QTL was found on SSC7. In the same region of SSC7, a maternally expressed QTL affecting muscle depth was found. Chromosome 6 harbored a maternally expressed QTL on the short arm and a paternally expressed QTL on the long arm, both affecting intramuscular fat content. The individual QTL explained from 2% up to 10% of the phenotypic variance. The known homologies to human and mouse did not reveal positional candidate genes. This study demonstrates that testing for imprinting should become a standard procedure to unravel the genetic control of multifactorial traits.

title: Detection and characterization of quantitative trait loci for growth and reproduction traits in pigs
abstract: A genome scan was performed on F 2 animals of a cross between Meishan and Dutch commercial pigs. Phenotypic data were available for growth traits and ultrasonic backfat thickness on 942–1151 animals, and for litter size on 249 and 206 animals at first and second parity, respectively. QTL analyses were performed using interval mapping by regression under the line-cross approach complemented by tests for genomic imprinting and sex-specific QTL effects. For backfat thickness, the analyses revealed significant QTL on chromosomes 2, 7, 14, and X, with significant imprinting for chromosomes 2 and 14. Significant QTL were detected for the different growth traits on chromosomes 1, 4, 7, and 8. Both the QTL on chromosome 4 and chromosome 8 showed maternal expression for a specific growth stage. The QTL analyses for litter size revealed one suggestive QTL for first parity and three suggestive QTL for the second parity. Analyses under a half-sib model did not reveal additional significant QTL, but confirmed several of the QTL detected under the line-cross models. This study provides confirmation of several QTL affecting growth and fat deposition in pigs and adds interesting new insight into their mode of expression. Furthermore, additional significant and suggestive QTL were identified.

title: Multiple trait analysis of genetic mapping for quantitative trait loci.
abstract: We present in this paper models and statistical methods for performing multiple trait analysis on mapping quantitative trait loci (QTL) based on the composite interval mapping method. By taking into account the correlated structure of multiple traits, this joint analysis has several advantages, compared with separate analyses, for mapping QTL, including the expected improvement on the statistical power of the test for QTL and on the precision of parameter estimation. Also this joint analysis provides formal procedures to test a number of biologically interesting hypotheses concerning the nature of genetic correlations between different traits. Among the testing procedures considered are those for joint mapping, pleiotropy, QTL by environment interaction, and pleiotropy vs. close linkage. The test of pleiotropy (one pleiotropic QTL at a genome position) vs. close linkage (multiple nearby nonpleiotropic QTL) can have important implications for our understanding of the nature of genetic correlations between different traits in certain regions of a genome and also for practical applications in animal and plant breeding because one of the major goals in breeding is to break unfavorable linkage. Results of extensive simulation studies are presented to illustrate various properties of the analyses.

title: Advanced intercross lines, an experimental population for fine genetic mapping.
abstract: An advanced intercrossed line (AIL) is an experimental population that can provide more accurate estimates of quantitative trait loci (QTL) map location than conventional mapping populations. An AIL is produced by randomly and sequentially intercrossing a population that initially originated from a cross between two inbred lines or some variant thereof. This provides increasing probability of recombination between any two loci. Consequently, the genetic length of the entire genome is stretched, providing increased mapping resolution. In this way, for example, with the same population size and QTL effect, a 95% confidence interval of QTL map location of 20 cM in the F2 is reduced fivefold after eight additional random mating generations (F10). Simulation results showed that to obtain the anticipated reduction in the confidence interval, breeding population size of the AIL in all generations should comprise an effective number of > or = 100 individuals. It is proposed that AILs derived from crosses between known inbred lines may be a useful resource for fine genetic mapping.