SHORT COMMUNICATION doi:10.1111/j.1365-2052.2004.01165.x Comparative mapping of human 10 to pig 10 and 14*

D. Nonneman and G. A. Rohrer USDA, ARS, US Meat Animal Research Center, Clay Center, NE, USA

Summary Identification of predictive markers in QTL regions that impact production traits in com- mercial populations of swine is dependent on construction of dense comparative maps with human and mouse genomes. Chromosomal painting in swine suggests that large genomic blocks are conserved between pig and human, while mapping of individual reveals that order can be quite divergent. High-resolution comparative maps in regions affecting traits of interest are necessary for selection of positional candidate genes to evaluate nucleotide variation causing phenotypic differences. The objective of this study was to construct an ordered comparative map of human and pig chro- mosomes 10 and 14. As a large portion of both pig chromosomes are represented by HSA10, genes at regularly spaced intervals along this chromosome were targeted for placement in the porcine genome. A total of 29 genes from human chromosome 10 were mapped to porcine chromosomes 10 (SSC10) and 14 (SSC14) averaging about 5 Mb dis- tance of human DNA per marker. Eighteen genes were assigned by linkage in the MARC mapping population, five genes were physically assigned with the IMpRH mapping panel and seven genes were assigned on both maps. Seventeen genes from human 10p mapped to SSC10, and 12 genes from human 10q mapped to SSC14. Comparative maps of mam- malian species indicate that chromosomal segments are conserved across several species and represent syntenic blocks with distinct breakpoints. Development of comparative maps containing several species should reveal conserved syntenic blocks that will allow us to better define QTL regions in livestock.

Keywords comparative map, gene, human chromosome 10, microsatellite, SNP, SSC10, SSC14.

Sufficient variation in production traits exists in commercial quite divergent. High-resolution comparative maps in populations of livestock to exploit allelic variation of regions affecting traits of interest are necessary for devel- superior animals to increase production efficiency and opment of additional neighbouring markers and selection of improve the quality of livestock products. Identification of positional candidate genes to evaluate for nucleotide vari- predictive markers that impact production traits in swine ation causing phenotypic differences. Although high-den- can be achieved by constructing dense comparative maps sity physical and linkage maps exist in the pig, gene order with human and mouse genomes. Bidirectional chromoso- for most regions is poorly defined. The sequencing of vast mal painting experiments suggest that large genomic blocks numbers of porcine ESTs and SNP genotyping has greatly are conserved between pig and human (Johansson et al. enabled mapping and assignment of a large number of 1995; Goureau et al. 1996), while mapping of individual genes to comparative maps (Fahrenkrug et al. 2002a). genes or regions suggest that for many regions gene order is The objective of this study was to construct a systematic ordered comparative map of human chromosome 10 with pig chromosomes 10 and 14. Several economically Address for correspondence important reproductive and carcass traits have been dis-

D. Nonneman, USDA, ARS, US Meat Animal Research Center, Spur covered on both pig chromosomes (Rohrer & Keele 1998a,b; 18D, PO Box 166, Clay Center, NE 68933-0166, USA. Rohrer et al. 1999; Rohrer 2000; Wada et al. 2000; Hirooka et al. 2001; Malek et al. 2001a,b). As a large por- E-mail: [email protected] *The nucleotide sequence data reported in this paper have been sub- tion of both pig chromosomes are represented by HSA10, mitted to GenBank and have been assigned the accession numbers genes at regularly spaced intervals along this chromosome AY368174–AY368203. with available porcine sequence were targeted for place- Accepted for publication 2 June 2004 ment in the porcine genome.

338 2004 International Society for Animal Genetics, Animal Genetics, 35, 338–343 Comparative mapping of porcine chromosomes 10 and 14 339

Primer pairs for amplification of genomic DNA were assigned with the IMpRH mapping panel and seven genes designed from porcine EST sequences in TIGR Gene Indices were assigned on both maps. Seventeen genes from human (TIGR website; http://www.tigr.org/tdb/ssgi/) or from 10p mapped to SSC10, and 12 genes from human 10q sequences deposited in GenBank (Table 1) using Primer 3 mapped to SSC14. (Rozen & Skaletsky 2000; code available at: http://www- Seventeen loci from human chromosome 10p all mapped genome.wi.mit.edu/genome_software/other/primer3.html). to SSC10q (Table 1). This corresponds to about 30 Mb of Genes were targeted by their position on human chromo- human DNA that is represented by about 60 cM on the some 10 (UCSC version hg16 based on NCBI Build 34; porcine linkage map or an average distance of <4 cM Golden Path website; http://genome.ucsc.edu/) and avail- (>2 Mb of human DNA) per marker. Thirteen genes were ability of porcine EST sequence. Intronic and 3¢-UTR regions linkage mapped using SNP, 10 loci were mapped by ra- of porcine genes were targeted for amplification and diation hybrid mapping (Table 2); seven loci (AKR1C2, sequencing for SNP discovery in a PTC-200 DNA engine CREM, EPC1, GDI2, IDI1, MRC1 and VIM) were placed on (MJ Research Inc, Watertown, MA, USA) using Hot Star both maps. Three genes (GATA3, NUDT5 and SUV39H2) Taq polymerase (Qiagen, Valencia, CA, USA) and 100 ng of were mapped using the RH panel, because the SNP found in genomic DNA in 25 ll reactions (Fahrenkrug et al. 2002b). the sequence were not informative enough to map them on Amplicons that contained a SNP that was heterozygous the linkage map (Table 2). CREM was mapped using two in the common sire or in at least two of the dams were microsatellites (SB79 and SB81) that were found in a BAC mapped using a primer extension assay on the Sequenom that contained CREM and on the RH panel because the MassArrayTM system (San Diego, CA, USA). Linkage ana- sequence did not contain SNP and the introns are too large lyses were performed for SNP across seven families to amplify across. The gene order is primarily arranged (86 progeny) of the MARC swine reference population from telomere to centromere of human 10p to the telomere (Rohrer et al. 1996; CRIMAP v2.4; Green et al. 1990). and centromere of porcine 10q, as previously described Multipoint locations for all mapped markers are based on (Hiraiwa et al. 2001; Nonneman & Rohrer 2003). On the latest published swine genetic map (Rohrer et al. 1996; SSC10, gene order is conserved from 105 to 127 cM, cor- http://www.marc.usda.gov/). responding to 5–13 Mb on human chromosome 10. Rear- Genes were physically mapped using the 118-clone rangement of genes near the centromere of SSC10q was INRA–University of Minnesota porcine 7000Rad Radiation found involving CREM, EPC1 and GAD2; PIP5K2A and Hybrid (IMpRH) panel (Yerle et al. 1998). Primers used BMI1. CREM is the most centromeric human 10p marker were described above, or designed from intron sequences that mapped near the center of SSC10q. Despite a com- obtained with the original EST primers. Amplifications were parative mapping approach targeting human genes, a large performed in 15 ll PCR using 12.5 ng panel DNA, 1.5 mM gap in marker coverage exists in the pig between 109 and MgCl2, 200 lM dNTPs, 1 lM each primer, 0.25 U Hot Star 125 cM (CUGBP2 and ITIH2; Fig. 1). This is a region Taq and 1X of supplied buffer. The PCR mixture was held at which corresponds to approximately 8–11 Mb on human 94 C for 15 min, and cycled 40 times at 94 C for 20 s, the 10p, an area devoid of any known genes or ESTs. This indicated annealing temperature for 30 s (Table 2) and could be a recombination hotspot in the pig, which would extension at 72 C for 1–1.5 min, followed by a final expand the linkage map in that region. No recombination extension at 72 C for 5 min. Data were analysed for two- was seen for two groups of markers, VIM, MRC1 and EPC1, point and multipoint linkage (Hawken et al. 1999) with the and for PIP5K2A and BMI1. However, the order for VIM, IMpRH mapping tool and submitted to the IMpRH database MRC1 and EPC1 was determined on the IMpRH panel (http://imprh.toulouse.inra.fr/). Carthagene (http:// (Table 2). www.inra.fr/bia/T/CarthaGene/) was used to estimate Twelve genes from human 10q were mapped to SSC14q; multipoint marker distance and order using all public DEPP was the only mapped using only the RH panel markers in the IMpRH database (http://imprh.tou and 11 genes were linkage mapped using SNP (Table 1). louse.inra.fr/) and those developed in this study to PPYR1 and NCOA were mapped using multiple SNP in the approximate position of RH mapped markers on the MARC same amplicon, and an indel was used to map CTBP2. DEPP linkage map. is the most centromeric human 10q marker that was A total of 29 genes from human chromosome 10 were assigned to the RH map. These markers cover about 60 cM mapped to porcine chromosomes 10 (SSC10) and 14 on the SSC14 and about 90 Mb of human DNA for an (SSC14) averaging about 5 Mb distance of human DNA per average marker distance of 5 cM in the pig or about 7.5 Mb marker. All amplicons were sequenced and had significant in human. Gene order was colinear for most of human 10q identity to the porcine and human target exon sequences. and SSC14q, except near the centromere where rear- These sequences have been entered into GenBank under rangements were found in the positions of CDC2 and the accession numbers AY368174–AY368203 and SFTPA1 (Fig. 2). No recombination was found between two AY515260. Eighteen genes were assigned by linkage in the groups of markers, MAPK8, SFTPA1 and PPYR1, and MARC mapping population, five genes were physically DLG5, NCOA4 and MSMB.

2004 International Society for Animal Genetics, Animal Genetics, ‘No claim to original US government works’, 35, 338–343 340 onmnadRohrer and Nonneman

Table 1 Genes from human chromosome 10 linkage mapped to porcine chromosomes 10 and 14.

Marker Gene name Forward primer Reverse primer Size Accession no. Polymorphism SSC (chr:cM) Hsa10 (Mb)

IDI-1 Isopentyl-diphosphate delta isomerase agataaacacggaccatctgga tcgttgttggagcagaagttta 450 AY368174 gctccYgacct ccttRccccg 10:126 1.0 GDI-2 GDP dissociation inhibitor 2 ctggagctcctggagccgattg tgatattaggaggggagggtct 550 AY368175 cccccScctcc 10:126 6.0 04ItrainlSceyfrAia Genetics, Animal for Society International 2004 GATA3 GATA binding factor-3 attaacagacccctgaccatga aagcctatatccatggggtttc 400 AY368199 None 10:126 8.3 CUGBP2 CUG triplet repeat, RNA binding 2 tctaaaggaaaaagctgcaagg gctcaaattccatcaaacaaca 500 AY368194 tagtcRttttt 10:109 11.5 SEC61A2 sec61 homologue tcgaggatcctgtccacgtgg ctcgtgaaccatggaggtgtc 610 AY368178 tacttYttcat 10:105 12.3 NUDT5 Nudix-type motif 5 ggagctcgaggaagagactggc gcctcacgttctcagcgtcatc 440 AY368203 None 10:103 12.4 CKLiK CAMKI-like protein kinase catccacgagtccgtcagcgc gaggctgcccgatacacttgc 900 AY368181 aaccaRcRtgg 10:107 12.5 MCM10 Minichromosome maintenance deficient 10 agaaacctcaggatgtcctgg cctggcaggctccagttcatc 1100 AY368180 cgggaRctcct 10:87 13.4 SUV39H2 Suppressor of variegation 3-9 homologue 2 gttgttcatgcacagattgctt tgtgtgcctttttgtacaatcc 420 AY368198 None 10:87 15.1 NMT2 Glycylpeptide N-tetradecanoyltransferase 2 agatgatgacaacatgttccg tttgctgggatggcgctgatg 300 AY368179 ggggcYtgggg tcaccYgcgtt 10:86 15.3 VIM Vimentin tgcaggatgagattcagaacat ttcagggaagaaaaatttggaa 600 AY368193 cagttYagaga ttatgMcttRg 10:79 17.4 MRC1 Mannose receptor C1 gataaggccaaacaccctca ccttgtctttgatgtttatttcaca 390 AY368183 gtaagRggtac 10:79 18.0 BMI-1 B-lymphoma Mo-MLV insertion region ggctctaatgaagatagaggag tcacagtcattgctgctgggca 660 AY368176 gtattWtatgc 10:91 22.0 NMA Putative agctgagcgcctgcttct ggtcagcttgcctcatcttatc 550 AY515260 None 10:75 29.0 MAP3K8 Mitogen-activated protein kinase kinase kinase tgcttcaaacatttaaaacgtga tggacaaacgtatggttttcat 300 AY368177 tcttaYgactc ttaaa[g]tatgt 10:72 30.9 nmlGenetics Animal EPC1 Enhancer of polycomb 1 tagtatgaggtggctttgggaat ccgacgcctctacagaaacata 520 AY368182 gtttaYctctc 10:79 32.7 CREM cAMP responsive element modulator agcagtgagaagctttggtcat ccactaaatggtgagctatcataa 360 AY368200 None 10:99 35.6 SB79 cAMP responsive element modulator aggtagctcctgatgtgaaagc gtagaaaagggagaggcatggt 153 AY368202 Microsatellite 10:98 35.6 SB81 cAMP responsive element modulator aagcaacaaaatgctctatgact gcttaacacagggtagggtgat 188 AY368192 Microsatellite 10:100 35.6

N li ooiia Sgvrmn works’, government US original to claim ‘No , DEPP Decidual protein induced by progesterone tcagctacagtttgcagtttcc tgacaaatggacctcacgtagt 180 AY368201 None 14:47 45.2 PPYR1 Neuropeptide Y receptor Y4 tataccaccttcctgctgctct tgctgagtgaccagaatgaact 770 AY368186 ctgtcYacagt ttttcRttgcc 14:45 46.9 MSMB Beta-microseminoprotein ccacctttgtgactttatgcaa gccacaagtacactcctcacag 550 AY368185 cccacYgagcg 14:48 51.4 NCOA4 Nuclear receptor coactivator 4 agattcacagctgcataagtcg gcgtagagcagttgtgtcagtt 1100 AY368184 tcataStaggt caccaSaKcca 14:47 51.5 CDC2 Cell division cycle 2 protein gccgaactagcaactaagaacc tttcatccaagtttttgacgtg 310 AY368188 RtaagYtgtta 14:41 62.4 DLG5 Drosophila large discs ggaggaggtctatgtggagatg tgtcatcaacatagaggatgtcg 1100 AY368190 gtactWgtggc 14:47 79.5 SFTPA1 Surfactant, pulmonary-associated protein A1 gcactccatgagatcagacatc gctctggcacacaactctctaa 600 AY368191 ccccaRgttcc 14:45 81.3 SCD Stearoyl-CoA desaturase cctgtggagtcaccgaactt gtggggatcagcatctgttt 1475 AY368187 cactaYtttag 14:54 102.2 NDUFB8 NADH dehydrogenase 1 beta subcomplex, 8 gtatggtgactacccgaagctc aaccgaagaggtgcttacacat 660 AY368196 gattaKYaaag 14:54 102.4 RGS10 Regulator of G-protein signalling 10 caacctcatgaagtatgacagc ggcctcacatgaagtttaatca 350 AY368195 tccccScaaag 14:73 121.4 CTBP2 C-terminal binding protein 2 caacgagcaatagcagagagtg tccccatctcgtaggacttaaa 490 AY368189 agtgt[t]aaagc 14:90 126.7 ECHS1 Enoyl-CoA hydratase ggagatggtccttactggtgac gccattgctgtgacaatcttag 570 AY368197 cacgcRcacag 14:104 135.1 35 338–343 , Comparative mapping of porcine chromosomes 10 and 14 341

Table 2 Markers and primers physically mapped on the 118-clone IMpRH 7000Rad panel.

Marker Forward primer Reverse primer Size Tm Chr Linked marker Dist. (R) % Ret. LOD

CREM agcagtgagaagctttggtcat ccactaaatggtgagctatcataa 360 60 10 SW305 0.11 40 22.3 AKR1C2 atcatcacctagggtcaacaattc gagctcccgctcagtactcttcag 350 55 10 SWR67 0.64 29 6.09 SUV39H2 gttgttcatgcacagattgctt tgtgtgcctttttgtacaatcc 200 58 10 SW920 0.24 45 15.07 DEPP tcagctacagtttgcagtttcc tgacaaatggacctcacgtagt 180 58 14 SW2593 0.23 29 13.95 DLG5 ggaggaggtctatgtggagatg tgtcatcaacatagaggatgtcg 1100 57 14 SW1536 0.72 18 4.68 GATA3 attaacagacccctgaccatga aagcctatatccatggggtttc 400 58 10 SW305 0.58 30 6.84 NUDT5 ggagctcgaggaagagactggc gcctcacgttctcagcgtcatc 440 55 10 SW1626 0.25 29 15.19 EPC1 tagtatgaggtggctttgggaat ccgacgcctctacagaaacata 520 58 10 SW920 0.45 32 8.46 MRC1 gataaggccaaacaccctca ccttgtctttgatgtttatttcaca 390 58 10 SW1991 0.56 31 7.27 VIM gatggctctcgacattgagatt ttggttttcacagcatcatagg 300 57 10 SW1991 0.44 34 9.44 NMA agctgagcgcctgcttct ggtcagcttgcctcatcttatc 1050 57 10 SW1041 0.42 25 9.64 GDI-2 ctggagctcctggagccgattg tgatattaggaggggagggtct 600 57 10 SWR67 0.43 28 9.23 IDI-1 gtcacctgaacgagaacatcaa tcgttgttggagcagaagttta 350 57 10 UMNp1176 0.24 33 12.8

HSA10 SSC14 HSA10 SSC10 0 0

40 CDC2 IMpRH MARC Linkage MMU18 MAPK8* 40 20 20 0 SSC25A02 MAP3K8 MMU8 SFTPA1 SSC10G07 NMA 100 MMU18 SWR158 EPC1 PPYR1 SWR334 LEA 200 MRC1 30 MMU18 NMA 60 VIM BFT OR 300 SW1991 MMU18 DEPP EPC1 NMT2 40 40 400 VIM SUV39H2 DLG5 MCM10 MMU6 500 MRC1 NCOA4 SW920 GAD2* NN 600 SUV39H2 80 PIP5K2A* MSMB MMU14 BMI1 20 MMU2 0 60 60 CREM GLUD1* BFT CREM CKLiK FSH 100 SW305 APT1* MMU10 SW951 100 NUDT5 200 SEC61A RBP4* SW1626 CUGBP2 NUDT5 80 NDUFB8 80 300 ITIH2* 10 MMU2 MMU14 GATA3 GATA3 SCD 400 SWR67 ATP5C1 * MMU14 GDI2 120 PRKCQ* NPM3* 500 AKR1C2 GDI2 SE77660* AP IDI1 AKR1C2* IDI1 MMU13 600 100 RGS10 100 cM 0 CTBP2 MMU19 cR Mb cM ECHS1 120 Figure 1 Comparative maps of chromosomes human 10p and porcine MMU7 10q. Maps shown are (left to right) partial pig RH map of 10q, MARC 140 linkage map of pig 10q and human 10p. Distances are in centirays (cR), Mb centimorgans (cM) and megabases (Mb), respectively. New markers are shown in bold type and boxes indicate no recombination between Figure 2 Comparative maps of chromosomes human 10q and porcine markers. Dotted lines indicate estimated position of RH markers on the 14q. New markers are shown in bold type and boxes indicate no porcine linkage map and asterisks indicate markers that were previously recombination between markers. Dotted lines indicate estimated reported (http://www.marc.usda.gov/). QTL positions are shown as position of RH markers on the porcine linkage map and asterisks bars on the left and the peak indicated with a symbol on the bar and on indicate markers that were previously reported (http://www.marc. the linkage map [OR (v), ovulation rate; NN (h), number of nipples; usda.gov/). QTL positions are shown as bars on the left with the peak BFT (x), backfat; FSH ()), plasma follicle-stimulating hormone of indicated with a symbol on the bar and on the linkage map [LEA ()), boars; AP (q), age at puberty of gilts] and are taken from references: loin eye area; BFT (h), backfat] taken from reference Rohrer & Keele Rohrer et al. 1999; Rohrer 2000; and Rohrer et al. 2001. Mouse 1998a,b. Mouse synteny is shown at right and is taken from the UCSC synteny is shown at right and is taken from the UCSC Genome Browser Genome Browser (http://genome.ucsc.edu/). (http://genome.ucsc.edu/). Direction of mouse chromosome is shown by arrow. one large inversion spans over 10 Mb of human chromo- some 10 (17–27 Mb), which corresponds to a region of The genetic mapping of genes in these syntenic groups is mouse chromosome 2 that is inverted in the mouse genome consistent with previously reported physical map results (http://genome.ucsc.edu/cgi-bin/hg), relative to human. (Hiraiwa et al. 2001; Rink et al. 2002; Wimmers et al. Another rearrangement on SSC10 that includes CREM, 2002; Cirera et al. 2003; Lahbib-Mansais et al. 2003; MAP3K8 and NMA (29–36 Mb) is a region corresponding Tuggle et al. 2003). Despite high levels of conserved synt- to mouse chromosome 18 that is rearranged. Rearrange- eny, differences in gene order between human and pig ment of gene order on SSC14 contains three regions span- usually correspond to regions where inversions or ning about 6 Mb each of human chromosome 10 (46–51, translocations in the murine genome occurred. On SSC10, 74–81 and 82–88 Mb) that are syntenic to mouse chro-

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mosome 14 which is fragmented and oriented in opposite Hirooka H., de Koning D.J., Harlizius B., van Arendonk J.A., Rattink directions. Another rearrangement in SSC14 includes A.P., Groenen M.A., Brascamp E.W. & Bovenhuis H. (2001) A CDC2, which is located on mouse chromosome 10. These whole-genome scan for quantitative trait loci affecting teat breakpoint regions are conserved in bovine chromosomes number in pigs. Journal of Animal Science 79, 2320–6. 13, 26 and 28, as well, and genes within these rearrange- Johansson M., Ellegren H. & Andersson L. (1995) Comparative mapping reveals extensive linkage conservation-but with gene ments are confined within conserved units or are distributed order rearrangements-between the pig and the human genomes. into fragmented regions among the three chromosomes Genomics 25, 682–90. (Larkin et al. 2003). Lahbib-Mansais Y., Tosser-Klopp G., Leroux S., Cabau C., Karsenty Comparative maps of mammalian species indicate that E., Milan D., Barillot E., Yerle M., Hatey F. & Gellin J. (2003) chromosomal segments are conserved across several species Contribution to high-resolution mapping in pigs with 101 type I (Larkin et al. 2003; Murphy et al. 2003) and represent markers and progress in comparative map between humans and syntenic blocks derived from intact ancestral chromosomes pigs. Mammalian Genome 14, 275–88. fissioned at distinct breakpoints. Development of higher Larkin D.M., Everts-van der Wind A., Rebeiz M. et al. (2003) A resolution comparative maps with several species should cattle-human comparative map built with cattle BAC-ends and reveal conserved syntenic blocks that will allow us to better sequence. Genome Research 13, 1966–72. define QTL regions in livestock. Malek M., Dekkers J.C., Lee H.K., Baas T.J. & Rothschild M.F. (2001a) A molecular genome scan analysis to identify chromo- somal regions influencing economic traits in the pig. I. Growth Acknowledgements and body composition. Mammalian Genome 12, 630–6. Malek M., Dekkers J.C., Lee H.K., Baas T.J., Prusa K. Huff-Lonergan The authors wish to thank the expert technical assistance of E., & Rothschild M.F. (2001b) A molecular genome scan analysis Bree Quigley for sequencing and cloning, Kris Simmerman to identify chromosomal regions influencing economic traits in for collection of microsatellite data, Linda Flathman for the pig. II. Meat and muscle composition. Mammalian Genome 12, mass spectrometry and Sherry Kluver for manuscript pre- 637–45. paration. We would also like to thank Dr Mark Rutherford Murphy W.J., Fronicke L., O’Brien S.J. & Stanyon R. (2003) The for supplying the IMpRH panel and Dr Denis Milan for origin of human chromosome 1 and its homologs in placental assistance with the IMpRH database. Mention of trade mammals. Genome Research 13, 1880–8. names or commercial products is solely for the purpose of Nonneman D.J. & Rohrer G.A. (2003) Comparative mapping of a region on chromosome 10 containing QTL for reproduction in providing information and does not imply recommendation, swine. Animal Genetics 34, 42–6. endorsement or exclusion of other suitable products by the Rink A., Santschi E.M., Eyer K.M., Roelofs B., Hess M., Godfrey M., US Department of Agriculture. Karajusuf E.K., Yerle M., Milan D. & Beattie C.W. (2002) A first- generation EST RH comparative map of the porcine and human References genome. Mammalian Genome 13, 578–87. Rohrer G.A. (2000) Identification of quantitative trait loci affecting Cirera S., Jorgensen C.B., Sawera M., Raudsepp T., Chowdhary B.P. & birth characters and accumulation of backfat and weight in a Fredholm M. (2003) Comparative mapping in the pig: localization Meishan-White Composite resource population. Journal of Animal of 214 expressed sequence tags. Mammalian Genome 14, 405–26. Science 78, 2547–53. Fahrenkrug S.C., Smith T.P., Freking B.A. et al. (2002a) Porcine Rohrer G.A. & Keele J.W. (1998a) Identification of quantitative trait gene discovery by normalized cDNA-library sequencing and EST loci affecting carcass composition in swine: I. Fat deposition cluster assembly. Mammalian Genome 13, 475–8. traits. Journal of Animal Science 76, 2247–54. Fahrenkrug S.C., Freking B.A., Smith T.P., Rohrer G.A. & Keele J.W. Rohrer G.A. & Keele J.W. (1998b) Identification of quantitative trait (2002b) Single nucleotide polymorphism (SNP) discovery in loci affecting carcass composition in swine: II. Muscling and porcine expressed genes. Animal Genetics 33, 186–95. wholesale product yield traits. Journal of Animal Science 76, 2255– Goureau A., Yerle M., Schmitz A., Riquet J., Milan D., Pinton P., 62. Frelat G. & Gellin J. (1996) Human and porcine correspondence Rohrer G.A., Alexander L.J., Hu Z., Smith T.P., Keele J.W. & Beattie, of chromosome segments using bi-directional chromosome C.W. (1996) A comprehensive map of the porcine genome. painting. Genomics 36, 252–62. Genome Research 6, 371–91. Green P., Falls K. & Crooks S. (1990) Documentation for CRIMAP, ver Rohrer G.A., Ford J.J., Wise T.H., Vallet J.L. & Christenson RK (1999) 2.4. Washington Univ. School of Medicine, St Louis, MO. Identification of quantitative trait loci affecting female reproductive Hawken R.J., Murtaugh J., Flickinger G.H., Yerle M., Robic A., traits in a multigeneration Meishan-White composite swine po- Milan D., Gellin J., Beattie C.W., Schook L.B. & Alexander L.J. pulation. Journal of Animal Science 77, 1385–1391. (1999) A first-generation porcine whole-genome radiation hybrid Rohrer G.A., Wise T.H., Lunstra D.D. & Ford J.J. (2001) Identifi- map. Mammalian Genome 10, 824–30. cation of genomic regions controlling plasma FSH concentrations Hiraiwa H., Uenishi H. & Yasue H. (2001) Alignment of VIM, in Meishan-White Composite boars. Physiological Genomics 6, MRC1, GAD2, and IL2RA genes on swine chromosome 10q by in 145–51. situ hybridization and RH mapping. Cytogenetics & Cell Genetics Rozen S. & Skaletsky H.J (2000) Primer3 on the WWW for general 94, 229–32. users and for biologist programmers. In: Bioinformatics Methods

2004 International Society for Animal Genetics, Animal Genetics, ‘No claim to original US government works’, 35, 338–343 Comparative mapping of porcine chromosomes 10 and 14 343

and Protocols: Methods in Molecular Biology (Ed. by S. Krawetz and Wimmers K., Ponsuksili S., Blaser U., Gellin J. & Schellander K. S. Misener), pp. 365–86. Humana Press, Totowa, NJ. (2002) Chromosomal assignments for porcine genes encoding Tuggle C.K., Green J.A., Fitzsimmons C. et al. (2003) EST-based in hepatic metabolic pathways. Animal Genetics 33, gene discovery in pig: virtual expression patterns and compar- 255–63. ative mapping to human. Mammalian Genome 14, 565–79. Yerle M., Pinton P., Robic A. et al. (1998) Construction of a whole- Wada Y., Akita T., Awata T. et al. (2000) Quantitative trait loci genome radiation hybrid panel for high-resolution gene mapping (QTL) analysis in a Meishan · Gottingen cross population. Ani- in pigs. Cytogenetics & Cell Genetics 82, 182–8. mal Genetics 31, 376–84.

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