Chromosome Research

, Volume 16, Issue 1, pp 109–127 | Cite as

The Horse Genome Derby: racing from map to whole genome sequence

  • Bhanu P. Chowdhary
  • Terje Raudsepp


The map of the horse genome has undergone unprecedented expansion during the past six years. Beginning from a modest collection of ∼300 mapped markers scattered on the 31 pairs of autosomes and the X chromosome in 2001, today the horse genome is among the best-mapped in domestic animals. Presently, high-resolution linearly ordered gene maps are available for all autosomes as well as the X and the Y chromosome. The approximately 4350 mapped markers distributed over the ∼2.68 Gbp long equine genome provide on average 1 marker every 620 kb. Among the most remarkable developments in equine genome analysis is the availability of the assembled sequence (EquCab2) of the female horse genome and the generation ∼1.5 million single nucleotide polymorphisms (SNPs) from diverse breeds. This has triggered the creation of new tools and resources like the 60K SNP-chip and whole genome expression microarrays that hold promise to study the equine genome and transcriptome in ways not previously envisaged. As a result of these developments it is anticipated that, during coming years, the genetics underlying important monogenic traits will be analyzed with improved accuracy and speed. Of larger interest will be the prospects of dissecting the genetic component of various complex/multigenic traits that are of vital significance for equine health and welfare. The number of investigations recently initiated to study a multitude of such traits hold promise for improved diagnostics, prevention and therapeutic approaches for horses.

Key words

Equus gene maps genome analysis horse 


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. Bailey E (1998) Odds on the FAST gene. Genome Res 8: 569–571.PubMedGoogle Scholar
  2. Bailey E, Binns MM (1998) The Horse Gene Map. ILAR J 39: 171–176.PubMedGoogle Scholar
  3. Barrey E, Mucher E, Robert C, Amiot F, Gidrol X (2006) Gene expression profiling in blood cells of endurance horses completing competition or disqualified due to metabolic disorder. Equine Vet J Suppl 36: 43–49.PubMedGoogle Scholar
  4. Beck J, Chowdhary BP, Brenig B (2005) Assignment of the equine colony stimulating factor 1 receptor gene (CSF1R) to equine chromosome 14q15→q16 (ECA14q15→q16) by in situ hybridization and radiation hybrid panel mapping. Cytogenet Genome Res 109: 533.PubMedGoogle Scholar
  5. Bellone R, Lear T, Durkin K, Adelson D, Bailey E (2005) FISH mapping Appaloosa candidate genes in the horse. 6th IEGMW, Dublin, Ireland.Google Scholar
  6. Boneker C, Kuiper H, Wohlke A, Drogemuller C, Chowdhary BP, Distl O (2005) Assignment of the COL16A1 gene to equine chromosome 2p15.1-p15.3 by FISH and confirmation by RH mapping. Anim Genet 36: 262–263.PubMedGoogle Scholar
  7. Boneker C, Kuiper H, Drogemuller C, Chowdhary BP, Distl O (2006) Molecular characterization of the equine collagen, type IX, alpha 2 (COL9A2) gene on horse chromosome 2p16→p15. Cytogenet Genome Res 115: 107–114.PubMedGoogle Scholar
  8. Brault LS, Famula TR, Penedo CM (2007) Association of equine cerebellar abiotrophy phenotype with markers on ECA2. 7th Dorothy Russell Havemeyer International Equine Genome Mapping Workshop, Tahoe City, CA, 37.Google Scholar
  9. Breen M, Lindgren G, Binns MM et al. (1997) Genetical and physical assignments of equine microsatellites—first integration of anchored markers in horse genome mapping. Mamm Genome 8: 267–273.PubMedGoogle Scholar
  10. Brenig B, Beck J, Hall AJ, Broad TE, Chowdhary BP, Piumi F (2004) Assignment of the equine solute carrier 26A2 gene (SLC26A2) to equine chromosome 14q15→q21 (ECA14q15→q21) by in situ hybridization and radiation hybrid panel mapping. Cytogenet Genome Res 107: 139.PubMedGoogle Scholar
  11. Bricker SJ, Brault LS, DelValle A, Millon LV, Murray JD, Penedo MC (2005) Radiation hybrid and linkage mapping of six new type I markers in the horse. Anim Genet 36: 182–184.PubMedGoogle Scholar
  12. Brinkmeyer-Langford C, Raudsepp T, Lee EJ et al. (2005) A high-resolution physical map of equine homologs of HSA19 shows divergent evolution compared with other mammals. Mamm Genome 16: 631–649.PubMedGoogle Scholar
  13. Brinkmeyer-Langford C, Raudsepp T, Gustafson-Seabury A et al. (2007) Comparative characterization of a 5Mb region on horse chromosome 21. Plant & Animal Genome XV, San Diego, CA, W235.Google Scholar
  14. Brooks SA, Bailey E (2005) Exon skipping in the KIT gene causes a Sabino spotting pattern in horses. Mamm Genome 16: 893–902.PubMedGoogle Scholar
  15. Brooks SA, Terry RB, Bailey E (2002) A PCR-RFLP for KIT associated with tobiano spotting pattern in horses. Anim Genet 33: 301–303.PubMedGoogle Scholar
  16. Brooks S, Lear TL, Adelson DL, Bailey E (2007) Characterization of the breakpoints of a chromosomal inversion associated with the Tobiano white spotting pattern in horses. Plant & Animal Genome XV, San Diego, CA, P599.Google Scholar
  17. Brown JJ, Ollier WE, Thomson W et al. (2006) TNF-alpha SNP haplotype frequencies in equidae. Tissue Antigens 67: 377–382.PubMedGoogle Scholar
  18. Brunberg E, Andersson L, Cothran G, Sandberg K, Mikko S, Lindgren G (2006) A missense mutation in PMEL17 is associated with the Silver coat color in the horse. BMC Genetics 7: 46.PubMedGoogle Scholar
  19. Bugno M, Slota E (2007) Application of arm-specific painting probes of horse X chromosome for karyotype analysis in an infertile Hutsul mare with 64,XX/65,XX+Xp karyotype: case report. Acta Vet Hung 55: 309–314.PubMedGoogle Scholar
  20. Bugno M, Slota E, Koscielny M (2007a) Karyotype evaluation among young horse populations in Poland. Schweiz Arch Tierheilkd 149: 227–232.PubMedGoogle Scholar
  21. Bugno M, Slota E, Pienkowska-Schelling A, Schelling C (2007b) Detection of equine X chromosome mosaicism in a mare using an equine X whole chromosome painting probe (WCPP)—a case report. Acta Vet Hung 55: 207–212.PubMedGoogle Scholar
  22. Caetano AR, Shiue YL, Lyons LA et al. (1999) A comparative gene map of the horse (Equus caballus). Genome Res 9: 1239–1249.PubMedGoogle Scholar
  23. Carbone L, Nergadze SG, Magnani E et al. (2006) Evolutionary movement of centromeres in horse, donkey, and zebra. Genomics 87: 777–782.PubMedGoogle Scholar
  24. Chowdhary BP, Bailey E (2003) Equine genomics: galloping to new frontiers. Cytogenet Genome Res 102: 184–188.PubMedGoogle Scholar
  25. Chowdhary BP, Gustavsson I (1992) Some aspects of equine genome analysis. Proceedings of an Annual Meeting of the German Society of Genetics, 1–3 October, Hannover, Germany, pp. 1–13.Google Scholar
  26. Chowdhary BP, Raudsepp T (2000) Cytogenetics and physical gene maps. In: Bowling AT, Ruvinsky A, eds., The Genetics of the Horse. Wallingford: CABI, pp. 171–242.Google Scholar
  27. Chowdhary BP, Raudsepp T (2005) Mapping genomes at the chromosome level. In: Ruvinsky A, Marshall Graves JA, eds., Mammalian Genomics. Wallingford: CABI, pp. 23–66.Google Scholar
  28. Chowdhary BP, Raudsepp T (2006) The Horse Genome. In: Volff J-N, ed., Vertebrate Genomes. Basel: Karger, pp. 97–110.Google Scholar
  29. Chowdhary BP, Raudsepp T, Fronicke L, Scherthan H (1998) Emerging patterns of comparative genome organization in some mammalian species as revealed by Zoo-FISH. Genome Res 8: 577–589.PubMedGoogle Scholar
  30. Chowdhary BP, Raudsepp T, Honeycutt D et al. (2002) Construction of a 5000(rad) whole-genome radiation hybrid panel in the horse and generation of a comprehensive and comparative map for ECA11. Mamm Genome 13: 89–94.PubMedGoogle Scholar
  31. Chowdhary BP, Raudsepp T, Kata SR et al. (2003) The first-generation whole-genome radiation hybrid map in the horse identifies conserved segments in human and mouse genomes. Genome Res 13: 742–751.PubMedGoogle Scholar
  32. Cothran EG, Graves KT, Strong D, Juras R, Ennis RB (2005) Linkage disequilibrium mapping of degenerative suspensory ligament desmitis in the Peruvian Paso, possible association with ECA 14q. 6th IEGMW, Dublin, Ireland.Google Scholar
  33. Dierks C, Momke S, Drogemuller C, Leeb T, Chowdhary BP, Distl O (2006) A high-resolution comparative radiation hybrid map of equine chromosome 4q12-q22. Anim Genet 37: 513–517.PubMedGoogle Scholar
  34. Dranchak PK, Ekenstedt KJ, Valberg SJ, Chowdhary BP, Raudsepp T, Mickelson JR (2006a) Chromosomal assignments for the equine AMPK family genes. Anim Genet 37: 293–294.PubMedGoogle Scholar
  35. Dranchak PK, Valberg SJ, Onan GW et al. (2006b) Exclusion of linkage of the RYR1, CACNA1S, and ATP2A1 genes to recurrent exertional rhabdomyolysis in Thoroughbreds. Am J Vet Res 67: 1395–1400.PubMedGoogle Scholar
  36. Dranchak PK, Leiper FC, Valberg SJ et al. (2007) Biochemical and genetic evaluation of the role of AMP-activated protein kinase in polysaccharide storage myopathy in Quarter horses. Am J Vet Res 68: 1079–1084.PubMedGoogle Scholar
  37. Durkin K, Raudsepp T, Chowdhary BP (2005) Characterization of two structural aberrations in the horse by FISH with BAC clones. 14th North American Colloquium on Animal Cytogenetics and Gene Mapping, Puerto Vallarta, Jalisco, Mexico, 9.Google Scholar
  38. Durkin K, Raudsepp T, Chowdhary BP (2008) Precise demarcation of the breakpoint in a Thoroughbred stallion carrying a Eca5 and Eca16 translocation. Plant & Animal Genome XVI, San Diego, USA.Google Scholar
  39. Ellis NA, Tammen I, Forhead AJ, Thomson P, Raadsma HW (2005) Characterization of the equine angiotensin converting enzyme gene. 6th IEGMW, Dublin, Ireland.Google Scholar
  40. Farber CR, Medrano JF (2004) Identification of putative homology between horse microsatellite flanking sequences and cross-species ESTs, mRNAs and genomic sequences. Anim Genet 35: 28–33.PubMedGoogle Scholar
  41. Giese A, Jude R, Kuiper H et al. (2002) Molecular characterization of the equine AEG1 locus. Gene 292: 65–72.PubMedGoogle Scholar
  42. Godard S, Vaiman D, Oustry A et al. (1997) Characterization, genetic and physical mapping analysis of 36 horse plasmid and cosmid-derived microsatellites. Mamm Genome 8: 745–750.PubMedGoogle Scholar
  43. Godard S, Schibler L, Oustry A, Cribiu EP, Guerin G (1998) Construction of a horse BAC library and cytogenetical assignment of 20 type I and type II markers. Mamm Genome 9: 633–637.PubMedGoogle Scholar
  44. Godard S, Vaiman A, Schibler L et al. (2000) Cytogenetic localization of 44 new coding sequences in the horse. Mamm Genome 11: 1093–1097.PubMedGoogle Scholar
  45. Goh G, Raudsepp T, Durkin K et al. (2007) High-resolution gene maps of horse chromosomes 14 and 21: additional insights into evolution and rearrangements of HSA5 homologs in mammals. Genomics 89: 89–112.PubMedGoogle Scholar
  46. Gu W, Bertone AL (2004) Generation and performance of an equine-specific large-scale gene expression microarray. Am J Vet Res 65: 1664–1673.PubMedGoogle Scholar
  47. Guérin G, Bailey E, Bernoco D et al. (1999) Report of the International Equine Gene Mapping Workshop: male linkage map. Anim Genet 30: 341–354.PubMedGoogle Scholar
  48. Gustafson-Seabury A, Raudsepp T, Goh G et al. (2005) High-resolution RH map of horse chromosome 22 reveals a putative ancestral vertebrate chromosome. Genomics 85: 188–200.PubMedGoogle Scholar
  49. Gustafson-Seabury A, Raudsepp T, Goh G et al. (2007) An integrated high-resolution whole genome radiation hybrid (RH) and comparative map for the horse. 7th Dorothy Russell Havemeyer International Equine Genome Mapping Workshop, Tahoe City, CA, 19.Google Scholar
  50. Gustafson AL, Tallmadge RL, Ramlachan N et al. (2003) An ordered BAC contig map of the equine major histocompatibility complex. Cytogenet Genome Res 102: 189–195.PubMedGoogle Scholar
  51. Haase B, Rieder S, Obexer-Ruff G et al. (2007) Molecular characterization of depigmentation phenotypes in horses. 7th Dorothy Russell Havemeyer International Equine Genome Mapping Workshop, Tahoe City, CA, 36.Google Scholar
  52. Hamann H, Jude R, Sieme H et al. (2007) A polymorphism within the equine CRISP3 gene is associated with stallion fertility in Hanoverian warmblood horses. Anim Genet 38: 259–264.PubMedGoogle Scholar
  53. Hansen M, Knorr C, Hall AJ, Broad TE, Brenig B (2007) Sequence analysis of the equine SLC26A2 gene locus on chromosome 14q15→q21. Cytogenet Genome Res 118: 55–62.PubMedGoogle Scholar
  54. Hanzawa K, Lear TL, Piumi F, Bailey E (2002) Mapping of equine potassium chloride co-transporter (SLC12A4) and amino acid transporter (SLC7A10) and preliminary studies on associations between SNPs from SLC12A4, SLC7A10 and SLC7A9 and osmotic fragility of erythrocytes. Anim Genet 33: 455–459.PubMedGoogle Scholar
  55. Hasegawa T, Tozaki T, Hirota K-I, Sato F, Oki H (2005) Linkage mapping of susceptive loci for recurrent exertional rhabdomyolysis (RER) in Thoroughbred. 6th IEGMW, Dublin, Ireland.Google Scholar
  56. Ing NH, Laughlin AM, Varner DD et al. (2004) Gene expression in the spermatogenically inactive “dark” and maturing “light” testicular tissues of the prepubertal colt. J Androl 25: 535–544.PubMedGoogle Scholar
  57. (ISCNH) Bowling AT, Breen M, Chowdhary BP et al. (1997) International system for cytogenetic nomenclature of the domestic horse. Report of the Third International Committee for the Standardization of the domestic horse karyotype, Davis, CA, USA, 1996. Chromosome Res 5: 433–443.PubMedGoogle Scholar
  58. Johannessen MK, Skretting G, Roed KH (2005) The Norwegian Equine Osteochondrosis Project—genetic expression in growth cartilage. 6th IEGMW, Dublin, Ireland.Google Scholar
  59. Jost U, Klukowska-Rotzler J, Dolf G et al. (2007) A region on equine chromosome 13 is linked to recurrent airway obstruction in horses. Equine Vet J 39: 236–241.PubMedGoogle Scholar
  60. Klukowska-Rotzler J, Bugno M, Sander P et al. (2006a) Chromosomal assignment of the two candidate genes (EGFR, CLCA1) for equine recurrent airway obstruction (RAO) by FISH and RH mapping. Hereditas 143: 138–141.PubMedGoogle Scholar
  61. Klukowska-Rotzler J, Jost U, Schelling C et al. (2006b) Characterization and RH mapping of six gene-associated equine microsatellite markers. Anim Genet 37: 305–306.PubMedGoogle Scholar
  62. Lear TL, Brandon R, Piumi F et al. (2001) Mapping of 31 horse genes in BACs by FISH. Chromosome Res 9: 261–262.PubMedGoogle Scholar
  63. Lear TL, Lundquist J, Gnerre S, Wade C, Lindblad-Toh K (2007) Anchoring supercontigs by FISH. 7th Dorothy Russell Havemeyer International Equine Genome Mapping Workshop, Tahoe City, CA.Google Scholar
  64. Lee EJ, Raudsepp T, Kata SR et al. (2004) A 1.4-Mb interval RH map of horse chromosome 17 provides detailed comparison with human and mouse homologues. Genomics 83: 203–215.PubMedGoogle Scholar
  65. Leeb T, Bruhn O, Philipp U, Kuiper H et al. (2005) Assignment of the equine S100A7 gene (psoriasin 1) to chromosome 5p12→p13 by fluorescence in situ hybridization and radiation hybrid mapping. Cytogenet Genome Res 109: 533.PubMedGoogle Scholar
  66. Leeb T, Vogl C, Zhu B et al. (2006) A human–horse comparative map based on equine BAC end sequences. Genomics 87: 772–776.PubMedGoogle Scholar
  67. Lefebvre-Lavoie J, Lussier JG, Theoret CL (2005) Profiling of differentially expressed genes in wound margin biopsies of horses using suppression subtractive hybridization. Physiol Genomics 22: 157–170.PubMedGoogle Scholar
  68. Lieto LD, Cothran EG (2003) The epitheliogenesis imperfecta locus maps to equine chromosome 8 in American Saddlebred horses. Cytogenet Genome Res 102: 207–210.PubMedGoogle Scholar
  69. Lindgren G, Breen M, Godard S et al. (2001) Mapping of 13 horse genes by fluorescence in situ hybridization (FISH) and somatic cell hybrid analysis. Chromosome Res 9: 53–59.PubMedGoogle Scholar
  70. Lindgren G, Sandberg K, Persson H et al. (1998) A primary male autosomal linkage map of the horse genome. Genome Res 8: 951–966.PubMedGoogle Scholar
  71. Locke MM, Ruth LS, Millon LV, Penedo MC, Murray JD, Bowling AT (2001) The cream dilution gene, responsible for the palomino and buckskin coat colours, maps to horse chromosome 21. Anim Genet 32: 340–343.PubMedGoogle Scholar
  72. Locke MM, Penedo MC, Bricker SJ, Millon LV, Murray JD (2002) Linkage of the grey coat colour locus to microsatellites on horse chromosome 25. Anim Genet 33: 329–337.PubMedGoogle Scholar
  73. MacLeod JN (2007) Transcriptional profiling of equine articular chondrocytes. 7th Dorothy Russell Havemeyer International Equine Genome Mapping Workshop, Tahoe City, CA, 42.Google Scholar
  74. Mariat D, Oustry-Vaiman A, Cribiu EP, Raudsepp T, Chowdhary BP, Guerin G (2001) Isolation, characterization and FISH assignments of horse BAC clones containing type I and II markers. Cytogenet Cell Genet 92: 144–148.PubMedGoogle Scholar
  75. Mariat D, Taourit S, Guerin G (2003) A mutation in the MATP gene causes the cream coat colour in the horse. Genet Sel Evol 35: 119–133.PubMedGoogle Scholar
  76. Marklund L, Moller MJ, Sandberg K, Andersson L (1996) A missense mutation in the gene for melanocyte-stimulating hormone receptor (MC1R) is associated with the chestnut coat color in horses. Mamm Genome 7: 895–899.PubMedGoogle Scholar
  77. Marti E, Binns M (1998) Horse genome mapping: a new era in horse genetics? Equine Vet J 30: 13–17.PubMedCrossRefGoogle Scholar
  78. Marti E, Glowatzki-Mullis M-L, Curik I, Torsteinsdottir S, Binns MM (2005) Investigating the genetic background for insect bite hypersensitivity in Icelandic horses. 6th IEGMW, Dublin, Ireland.Google Scholar
  79. Mau C (2003) Genetische Lokalisierung und Molekulare Analyse von “Dominant Weiss (W)” einer homozygot letalen Mutation beim Pferd. Veterinary Thesis. Zurich, Switzerland: ETH.Google Scholar
  80. McCue ME, Valberg SJ, Byrne K, Miller MB, Wade C, Mickelson JR (2007) Identification of a glycogen synthase 1 mutation resulting in polysaccharide storage myopathy. 7th Dorothy Russell Havemeyer International Equine Genome Mapping Workshop, Tahoe City, CA, 14.Google Scholar
  81. McGivney B, Gu J, Eivers S, Katz L, Hill E (2007) Population and functional genomics investigations of performance associations in thoroughbred horses. 7th Dorothy Russell Havemeyer International Equine Genome Mapping Workshop, Tahoe City, CA, 15.Google Scholar
  82. Metallinos DL, Bowling AT, Rine J (1998) A missense mutation in the endothelin-B receptor gene is associated with Lethal White Foal Syndrome: an equine version of Hirschsprung disease. Mamm Genome 9: 426–431.PubMedGoogle Scholar
  83. Mickelson JR, Wagner ML, Goh G et al. (2004) Thirty-five new equine microsatellite loci assigned to genetic linkage and radiation hybrid maps. Anim Genet 35: 481–484.PubMedGoogle Scholar
  84. Milenkovic D, Oustry-Vaiman A, Lear TL et al. (2002) Cytogenetic localization of 136 genes in the horse: comparative mapping with the human genome. Mamm Genome 13: 524–534.PubMedGoogle Scholar
  85. Milenkovic D, Chaffaux S, Taourit S, Guerin G (2003) A mutation in the LAMC2 gene causes the Herlitz junctional epidermolysis bullosa (H-JEB) in two French draft horse breeds. Genet Sel Evol 35: 249–256.PubMedGoogle Scholar
  86. Momozawa Y, Takeuchi Y, Tozaki T et al. (2005) Sequence, detection of polymorphisms and radiation hybrid mapping of the equine catechol-O-methyltransferase gene. Anim Genet 36: 190.PubMedGoogle Scholar
  87. Momozawa Y, Takeuchi Y, Tozaki T et al. (2006) Polymorphism identification, RH mapping, and association analysis with the anxiety trait of the equine serotonin transporter (SLC6A4) gene. J Vet Med Sci 68: 619–621.PubMedGoogle Scholar
  88. Momozawa Y, Takeuchi Y, Tozaki T et al. (2007) SNP detection and radiation hybrid mapping in horses of nine candidate genes for temperament. Anim Genet 38: 81–83.PubMedGoogle Scholar
  89. Mucher E, Jayr L, Rossignol F, Amiot F, Gidrol X, Barrey E (2006) Gene expression profiling in equine muscle tissues using mouse cDNA microarrays. Equine Vet J Suppl 36: 359–364.PubMedGoogle Scholar
  90. Muller D, Kuiper H, Boneker C et al. (2005a) Assignment of BGLAP, BMP2, CHST4, SLC1A3, SLC4A1, SLC9A5 and SLC20A1 to equine chromosomes by FISH and confirmation by RH mapping. Anim Genet 36: 457–461.PubMedGoogle Scholar
  91. Muller D, Kuiper H, Boneker C et al. (2005b) Physical mapping of the PTHR1 gene to equine chromosome 16q21.2. Anim Genet 36: 282–284.PubMedGoogle Scholar
  92. Muller D, Kuiper H, Momke S et al. (2005c) Assignment of the COMP gene to equine chromosome 21q12-q14 by FISH and confirmation by RH mapping. Anim Genet 36: 277–279.PubMedGoogle Scholar
  93. Murphy BA, Lear TL, Adelson DL, Fitzgerald BP (2007) Chromosomal assignments and sequences for the equine core circadian clock genes. Anim Genet 38: 84–85.PubMedGoogle Scholar
  94. Musilova P, Kubickova S, Zrnova E, Horin P, Vahala J, Rubes J (2007) Karyotypic relationships among Equus grevyi, Equus burchelli and domestic horse defined using horse chromosome arm-specific probes. Chromosome Res 15: 807–813.PubMedGoogle Scholar
  95. Myka JL, Lear TL, Houck ML, Ryder OA, Bailey E (2003a) FISH analysis comparing genome organization in the domestic horse (Equus caballus) to that of the Mongolian wild horse (E. przewalskii). Cytogenet Genome Res 102: 222–225.PubMedGoogle Scholar
  96. Myka JL, Lear TL, Houck ML, Ryder OA, Bailey E (2003b) Homologous fission event(s) implicated for chromosomal polymorphisms among five species in the genus Equus. Cytogenet Genome Res 102: 217–221.PubMedGoogle Scholar
  97. Nixon AJ, Glaser KE, Sun Q, Well ME (2008) Development and characterization of a novel large-scale whole transcript oligonucleotide exon array for the horse. Plant & Animal Genome XVI, San Diego, CA.Google Scholar
  98. Nomura M, Hosaka Y, Kasashima Y et al. (2007) Active expression of matrix metalloproteinase-13 mRNA in the granulation tissue of equine superficial digital flexor tendinitis. J Vet Med Sci 69: 637–639.PubMedGoogle Scholar
  99. Oakenfull EA, Buckle VJ, Clegg JB (1993) Localization of the horse (Equus caballus) alpha-globin gene complex to chromosome 13 by fluorescence in situ hybridization. Cytogenet Cell Genet 62: 136–138.PubMedCrossRefGoogle Scholar
  100. Paria N, Pearks Wilkerson AJ, Murphy WJ, Chowdhary BP, Raudsepp T (2007) Targeting Y chromosome genes in horses. 7th Dorothy Russell Havemeyer International Equine Genome Mapping Workshop, Tahoe City, CA, 23.Google Scholar
  101. Penedo MC, Millon LV, Bernoco D et al. (2005) International Equine Gene Mapping Workshop Report: a comprehensive linkage map constructed with data from new markers and by merging four mapping resources. Cytogenet Genome Res 111: 5–15.PubMedGoogle Scholar
  102. Perrocheau M, Boutreux V, Chadi-Taourit S et al. (2005) Equine FISH mapping of 36 genes known to locate on human chromosome ends. Cytogenet Genome Res 111: 46–50.PubMedGoogle Scholar
  103. Perrocheau M, Boutreux V, Chadi S et al. (2006) Construction of a medium-density horse gene map. Anim Genet 37: 145–155.PubMedGoogle Scholar
  104. Pielberg G, Mikko S, Sandberg K, Andersson L (2005) Comparative linkage mapping of the Grey coat colour gene in horses. Anim Genet 36: 390–395.PubMedGoogle Scholar
  105. Ramery E, Closset R, Bureau F, Art T, Lekeux P (2007) Relevance of using a human microarray to study gene expression in heaves-affected horses. Vet J [Epub. ahead of print].Google Scholar
  106. Raudsepp T, Fronicke L, Scherthan H, Gustavsson I, Chowdhary BP (1996) Zoo-FISH delineates conserved chromosomal segments in horse and man. Chromosome Res 4: 218–225.PubMedGoogle Scholar
  107. Raudsepp T, Kijas J, Godard S, Guerin G, Andersson L, Chowdhary BP (1999) Comparison of horse chromosome 3 with donkey and human chromosomes by cross-species painting and heterologous FISH mapping. Mamm Genome 10: 277–282.PubMedGoogle Scholar
  108. Raudsepp T, Mariat D, Guerin G, Chowdhary BP (2001) Comparative FISH mapping of 32 loci reveals new homologous regions between donkey and horse karyotypes. Cytogenet Cell Genet 94: 180–185.PubMedGoogle Scholar
  109. Raudsepp T, Kata SR, Piumi F et al. (2002a) Conservation of gene order between horse and human X chromosomes as evidenced through radiation hybrid mapping. Genomics 79: 451–457.PubMedGoogle Scholar
  110. Raudsepp T, Lear TL, Chowdhary BP (2002b) Comparative mapping in equids: the asine X chromosome is rearranged compared to horse and Hartmann’s mountain zebra. Cytogenet Genome Res 96: 206–209.PubMedGoogle Scholar
  111. Raudsepp T, Lee EJ, Kata SR et al. (2004a) Exceptional conservation of horse-human gene order on X chromosome revealed by high-resolution radiation hybrid mapping. Proc Natl Acad Sci USA 101: 2386–2391.PubMedGoogle Scholar
  112. Raudsepp T, Santani A, Wallner B et al. (2004b) A detailed physical map of the horse Y chromosome. Proc Natl Acad Sci USA 101: 9321–9326.PubMedGoogle Scholar
  113. Raudsepp T, Gustafson-Seabury A, Wagner M, Durkin K, Mickelson J, Chowdhary BP (2007a) The second generation radiation hybrid map (RH II) of the horse genome. Plant & Animal Genome XV, San Diego, CA, W108.Google Scholar
  114. Raudsepp T, Paria N, Durkin K, Murphy WJ, Chowdhary BP (2007b) A detailed physical map of the horse Y chromosome. Plant & Animal Genome XV, San Diego, CA, W110.Google Scholar
  115. Reeben M, Koho NM, Raekallio M, Hyyppa S, Poso AR (2006) MCT1 and CD147 gene polymorphisms in standardbred horses. Equine Vet J Suppl 36: 322–325.PubMedGoogle Scholar
  116. Reissmann M, Bierwolf J, Brockmann GA (2007) Two SNPs in the SILV gene are associated with silver coat colour in ponies. Anim Genet 38: 1–6.PubMedGoogle Scholar
  117. Rieder S, Taourit S, Mariat D, Langlois B, Guerin G (2001) Mutations in the agouti (ASIP), the extension (MC1R), and the brown (TYRP1) loci and their association to coat color phenotypes in horses (Equus caballus). Mamm Genome 12: 450–455.PubMedGoogle Scholar
  118. Rios JJ, Perelygin AA, Long MT et al. (2007) Characterization of the equine 2–5 oligoadenylate synthetase 1 (OAS1) and ribonuclease L (RNASEL) innate immunity genes. BMC Genomics 8: 313.PubMedGoogle Scholar
  119. Rosengren Pielberg G, Golovko A, Sundstrom E et al. (2007) Positional identification of the grey coat color mutatio in horse. 7th Dorothy Russell Havemeyer International Equine Genome Mapping Workshop, Tahoe City, CA, 13.Google Scholar
  120. Rudolph JA, Spier SJ, Byrns G, Rojas CV, Bernoco D, Hoffman EP (1992) Periodic paralysis in quarter horses: a sodium channel mutation disseminated by selective breeding. Nat Genet 2: 144–147.PubMedGoogle Scholar
  121. Santschi EM, Purdy AK, Valberg SJ, Vrotsos PD, Kaese H, Mickelson JR (1998) Endothelin receptor B polymorphism associated with lethal white foal syndrome in horses. Mamm Genome 9: 306–309.PubMedGoogle Scholar
  122. Shin EK, Perryman LE, Meek K (1997) A kinase-negative mutation of DNA-PK(CS) in equine SCID results in defective coding and signal joint formation. J Immunol 158: 3565–3569.PubMedGoogle Scholar
  123. Smith KJ, Bertone AL, Weisbrode SE, Radmacher M (2006) Gross, histologic, and gene expression characteristics of osteoarthritic articular cartilage of the metacarpal condyle of horses. Am J Vet Res 67: 1299–1306.PubMedGoogle Scholar
  124. Solberg OD, Jackson KA, Millon LV et al. (2004) Genomic characterization of equine interleukin-4 receptor alpha-chain (IL4R). Vet Immunol Immunopathol 97: 187–194.PubMedGoogle Scholar
  125. Spirito F, Charlesworth A, Linder K, Ortonne JP, Baird J, Meneguzzi G (2002) Animal models for skin blistering conditions: absence of laminin 5 causes hereditary junctional mechanobullous disease in the Belgian horse. J Invest Dermatol 119: 684–691.PubMedGoogle Scholar
  126. Swinburne J, Gerstenberg C, Breen M et al. (2000) First comprehensive low-density horse linkage map based on two 3-generation, full-sibling, cross-bred horse reference families. Genomics 66: 123–134.PubMedGoogle Scholar
  127. Swinburne JE, Hopkins A, Binns MM (2002) Assignment of the horse grey coat colour gene to ECA25 using whole genome scanning. Anim Genet 33: 338–342.PubMedGoogle Scholar
  128. Swinburne JE, Boursnell M, Hill G et al. (2006) Single linkage group per chromosome genetic linkage map for the horse, based on two three-generation, full-sibling, crossbred horse reference families. Genomics 87: 1–29.PubMedGoogle Scholar
  129. Swinburne JE, Blott SC, Klukowska J, Temperton E, Vaudin M, Gerber V (2007) Mapping genes associated with recurrent airway obstruction in horses. 7th Dorothy Russell Havemeyer International Equine Genome Mapping Workshop, Tahoe City, CA, 16.Google Scholar
  130. Takahashi T, Yawata M, Raudsepp T et al. (2004) Natural killer cell receptors in the horse: evidence for the existence of multiple transcribed LY49 genes. Eur J Immunol 34: 773–784.PubMedGoogle Scholar
  131. Tallmadge RL, Lear TL, Antczak DF (2005) Genomic characterization of MHC class I genes of the horse. Immunogenetics 57: 763–774.PubMedGoogle Scholar
  132. Terry RB, Archer S, Brooks S, Bernoco D, Bailey E (2004) Assignment of the appaloosa coat colour gene (LP) to equine chromosome 1. Anim Genet 35: 134–137.PubMedGoogle Scholar
  133. Tozaki T, Swinburne J, Hirota K, Hasegawa T, Ishida N, Tobe T (2007) Improved resolution of the comparative horse-human map: investigating markers with in silico and linkage mapping approaches. Gene 392: 181–186.PubMedGoogle Scholar
  134. Trifonov V, Yang F, Ferguson-Smith MA, Robinson TJ (2003) Cross-species chromosome painting in the Perissodactyla: delimitation of homologous regions in Burchell's zebra (Equus burchellii) and the white (Ceratotherium simum) and black rhinoceros (Diceros bicornis). Cytogenet Genome Res 103: 104–110.PubMedGoogle Scholar
  135. Tryon RC, White SD, Bannasch DL (2007) Homozygosity mapping approach identifies a missense mutation in equine cyclophilin B (PPIB) associated with HERDA in the American Quarter Horse. Genomics 90: 93–102.PubMedGoogle Scholar
  136. Wade CM, Gnerre S, Garber M, Lander E, Lindblad-Toh K (2007) Preliminary analysis of the equine genome sequence. 7th Dorothy Russell Havemeyer International Equine Genome Mapping Workshop, Tahoe City, CA, 6.Google Scholar
  137. Wagner ML, Goh G, Wu JT et al. (2004a) Sixty-seven new equine microsatellite loci assigned to the equine radiation hybrid map. Anim Genet 35: 484–486.PubMedGoogle Scholar
  138. Wagner ML, Goh G, Wu JT et al. (2004b) Radiation hybrid mapping of 63 previously unreported equine microsatellite loci. Anim Genet 35: 159–162.PubMedGoogle Scholar
  139. Wagner ML, Goh G, Wu JT et al. (2004c) Radiation hybrid mapping of 75 previously unreported equine microsatellite loci. Anim Genet 35: 68–71.PubMedGoogle Scholar
  140. Wagner ML, Raudsepp T, Goh G et al. (2006) A 1.3-Mb interval map of equine homologs of HSA2. Cytogenet Genome Res 112: 227–234.PubMedGoogle Scholar
  141. Ward TL, Valberg SJ, Lear TL et al. (2003) Genetic mapping of GBE1 and its association with glycogen storage disease IV in American Quarter horses. Cytogenet Genome Res 102: 201–206.PubMedGoogle Scholar
  142. Wittwer C, Chowdhary BP, Distl O (2005) Radiation hybrid mapping of equine CDK2, DGKA, DNAJC14, MMP19, CTSL and GAS1. Anim Genet 36: 536–537.PubMedGoogle Scholar
  143. Woehlke A, Nordsiek G, Scharfe M et al. (2008) Development of a BAC-based physical map of the horse genome. Plant & Animal Genome XVI, 2008, San Diego, CA, USA.Google Scholar
  144. Yang F, Fu B, O’Brien PC, Robinson TJ, Ryder OA, Ferguson-Smith MA (2003) Karyotypic relationships of horses and zebras: results of cross-species chromosome painting. Cytogenet Genome Res 102: 235–243.PubMedGoogle Scholar
  145. Yang F, Fu B, O’Brien PC, Nie W, Ryder OA, Ferguson-Smith MA (2004) Refined genome-wide comparative map of the domestic horse, donkey and human based on cross-species chromosome painting: insight into the occasional fertility of mules. Chromosome Res 12: 65–76.PubMedGoogle Scholar
  146. Yang GC, Croaker D, Zhang AL, Manglick P, Cartmill T, Cass D (1998) A dinucleotide mutation in the endothelin-B receptor gene is associated with lethal white foal syndrome (LWFS); a horse variant of Hirschsprung disease. Hum Mol Genet 7: 1047–1052.PubMedGoogle Scholar
  147. Young AE, Bower LP, Affolter VK, De Cock HE, Ferraro GL, Bannasch DL (2007) Evaluation of FOXC2 as a candidate gene for chronic progressive lymphedema in draft horses. Vet J 174: 397–399.PubMedGoogle Scholar
  148. Zabek T, Bugno M, Klukowska-Rotzler J, Uhlmann B, Gerber V, Slota E (2007) Chromosomal assignment of five equine genes responsible for the development of the skeletal and nervous systems. Anim Genet 38: 425–426.PubMedGoogle Scholar

Copyright information

© Springer 2008

Authors and Affiliations

  1. 1.Department of Veterinary Integrative Biosciences, College of Veterinary Medicine and Biomedical SciencesTexas A&M UniversityCollege StationUSA

Personalised recommendations