261:, may be responsible for the most familiar expressions of white: heterozygotes possessing common-size "blankets" and homozygotes possessing extensive "blankets" that may affect the entire coat. Even horses with extensive white usually retain dark colored regions just above the hooves, on the knees and hocks, stifles and elbows, hips and points of shoulder, the tail, mane, and the bony parts of the face. The smallest amount of white patterning is just a sprinkling of white over the hips.
825:
577:
426:
486:
456:
22:
544:
244:
844:" in a cave in France depict spotted horses with a leopard pattern. Archaeologists had debated over whether the artists were painting what they saw or whether the spotted horses had some symbolic meaning. However, a 2011 study of the DNA of ancient horses found that leopard complex was present, and therefore the cave painters most likely did see real spotted horses.
471:
441:
559:
93:, the spotting and especially the white regions associated with the leopard complex tend to be symmetrical and originate over the hips. Furthermore, a certain amount of this inherited white patterning is present at birth. The amount of white, even if none is present at birth, often grows throughout the horse's life by gradual "roaning" which is not related to
1535:
411:
307:
512:
527:
270:
584:
Although the spotting and roaning patterns that make up the leopard complex sometimes appear very different from each other, the ability of leopard-spotted horses to produce the full spectrum of patterns, from mottled skin to roaning to more leopard-spotted offspring, has long suggested that a single
502:
A distinct version of the leopard complex. Intermixed dark and light hairs with lighter colored area on the forehead, jowls and frontal bones of the face, over the back, loin and hips. Darker areas may appear along the edges of the frontal bones of the face as well and also on the legs, stifle, above
289:
gene will progressively lose pigment in both the skin and hair as they age. Also unlike graying out, the leopard spots are not affected by this roaning process. Neither are the "bony prominences" strongly affected. As a varnish roan horse lightens, the leopard spots indistinguishable from the rest of
766:
in horses has many causes, including eye trauma, disease, and bacterial, parasitic and viral infections, but ERU is characterized by recurring episodes of uveitis, rather than a single incident. If not treated, ERU can lead to blindness, which occurs more often in
Appaloosas than in other breeds. Up
721:
Congenital stationary night blindness has been linked with the leopard complex since the 1970s. The presence of CSNB in non-leopard breeds and horses suggested that the two conditions might be located on close, but separate genes. However, one study used ERG findings to diagnose all the homozygous
256:
or white leg markings, obscure leopard spots. A certain amount of leopard-associated white patterning may be present at birth. Temporal changes in the amount of white patterning are discussed below. Leopard-associated white patterning is usually symmetrical and originates over the hips. A proposed
701:
in horses which is present at birth (congenital), non-progressive (stationary) and affects the animal's vision in conditions of low lighting. Horses with CSNB may be hesitant to enter dimly-lit places - such as indoor arenas, dark stalls, or trailers - and be apprehensive when in such conditions,
202:
The presence of regions of alternating pigmented and unpigmented skin may not definitively suggest the leopard gene. They may not be visible due to the effects of other genes. For example, extensive white markings on the face may mask the presence of mottling around the eyes and muzzle, and white
234:
gene, and directly reflect the underlying coat color (bay, black, gray, cremello, red dun, and so on). Since these spots match the coat color, they are not visible unless the surrounding pigment is removed. As a rule, heterozygous leopards have larger, more abundant spots, while homozygotes have
767:
to 80% of all uveitis cases are found in
Appaloosas, with physical characteristics including light colored coat patterns, little pigment around the eyelids and sparse hair in the mane and tail denoting more at-risk individuals. Researchers may have identified a gene region containing an
702:
which may interfere with handling or riding. CSNB is usually diagnosed based on the owner's observations, but some horses have visibly abnormal eyes: poorly aligned eyes (dorsomedial strabismus) or involuntary eye movement (nystagmus). The condition can be confirmed using
285:. Horses with coat patterns within the leopard complex are known for their mystifying coat changes. This unusual characteristic is due at least in part to leopard roaning, also called "varnish roaning." While the gray gene only affects the hair, some horses with the
319:
horses with extensive white patterning at birth are white with large, self-colored spots. They are termed "leopard" if fully white, "near-leopard" if not. By the action of varnish roan, a near-leopard may in time become nearly indistinguishable from a full
251:
There is at least one genetically controlled type of white patterning that is strictly associated with the leopard complex. These white patterns permit the spots associated with the leopard complex to become visible. Other white patterns, such as
290:
the coat become visible. Some horses without any dense white patterning at birth seem to spontaneously develop into white, leopard-spotted horses with maturity. Varnishing is more common among
Appaloosa horses, and less common among Norikers and
183:
that is mottled, speckled or blotchy around the muzzle, eyes, genitals, and anus; the remainder of the body may be primarily pigmented (gray or black in the absence of other genes), primarily unpigmented (pink or flesh-colored), or
811:
has been recorded since ancient times, and suggests that the gene is very old. Leopard complex patterns may exist in low frequencies among some other breeds, depending on whether horses with leopard complex genetics existed in the
353:
horses with only a tiny amount of white patterning may not possess enough white to reveal large or small spots. A sprinkling of white patterning over the hips is called a "snowflake" pattern. Such tiny blankets may varnish and
302:
Like much of coat color genetics, commonly used terms do not necessarily correspond to precise genetic states. Nevertheless, terminology can reveal a lot about the genetic interactions surrounding the leopard complex.
1130:
Sandmeyer, Lynne S.; Breaux CB; Archer S; Grahn BH (2007). "Clinical and electroretinographic characteristics of congenital stationary night blindness in the
Appaloosa and the association with the leopard complex".
334:
horses with extensive white patterning at birth are white with tiny, sparse spots or none at all. In most languages, such foals are called "white-born" but the term familiar to most
English speakers is "fewspot
1375:
912:
714:, which transmit the nerve impulse to the next set of neurons. It is thought that these cells fail to undergo the basic chemical reaction for nerve impulse transmission, which involves shuttling of
327:
horses with less white patterning are described by the size of their "blanket" and the presence of spots: spotted blanket over loin and hips, for example. Again, these horses may varnish with age.
365:
horses, in the absence of dense white patterning, appear much the same. That is, unless they begin to varnish. As the coat becomes more and more white, spots may become visible. A homozygous
738:, a key factor in the transmission of nerve impulses. This protein, which is found in the retina and the skin, existed in fractional percentages of the normal levels in homozygous
230:
horses possess the aforementioned characteristics, heterozygotes and homozygotes differ significantly in the presence of true spots. True leopard spots are produced only by the
369:
horse, with only tiny spots, may simply develop this unique roaning pattern and is called "frosted" or "marble." A heterozygote may eventually show conspicuous leopard spots.
1074:"Differential Gene Expression of TRPM1, the Potential Cause of Congenital Stationary Night Blindness and Coat Spotting Patterns (LP) in the Appaloosa Horse (Equus caballus)"
378:
Base colors are overlain by various spotting patterns, which are variable and often do not fit neatly into a specific category. These patterns are described as follows:
247:
The extent of this horse's white patterning is illustrated because he is wet. Only large regions of pigmented (gray or black) skin will yield chestnut-colored hairs.
113:
dun-buckskin. A number of factors, each separately, genetically controlled, interact to produce familiar patterns such as "snowflake," "leopard," and "fewspot".
1379:
1476:
920:
503:
the eye, point of the hip and behind the elbow. The dark points over bony areas are called "varnish marks" and distinguish this pattern from a traditional
277:
Just as there is white patterning specifically associated with the leopard complex, there is a type of progressive roaning that is unrelated to
203:
markings on the legs often end in white hooves. Furthermore, other genes may produce similar conditions: white sclera are associated with broad
1289:"Evidence for a Retroviral Insertion in TRPM1 as the Cause of Congenital Stationary Night Blindness and Leopard Complex Spotting in the Horse"
89:
Coat patterns in the leopard complex range from being hardly distinguishable from an unaffected coat, to nearly pure white. Unlike most other
1021:
69:
skin. The leopard complex gene is also linked to abnormalities in the eyes and vision. These patterns are most closely identified with the
633:
140:; an incomplete dominant gene produces a different result depending on whether one or two copies are present. A horse's genotype may be
865:
Terry, RB; Archer S; Brooks S; Bernoco D; Bailey E (2004). "Assignment of the appaloosa coat colour gene (LP) to equine chromosome 1".
1469:
706:, from which a "negative ERG" indicates CSNB. While the retina is a normal shape, the nerve signal triggered when light reaches
1425:"Abstracts: 36th Annual Meeting of the American College of Veterinary Ophthalmologists, Nashville, TN, US, October 12–15, 2005"
451:
Considered an extension of a blanket to cover the whole body. A white horse with dark spots that flow out over the entire body.
481:
A horse with white spots, flecks, on a dark body. Typically the white spots increase in number and size as the horse ages.
310:
The reddish leopard-spots in this otherwise bay coat are only visible after leopard-related roaning has occurred, as here.
1462:
436:
a white blanket which has dark spots within the white. The spots are usually the same color as the horse's base color.
2005:
1986:
1955:
1945:
1920:
539:
Horses with roaning over the croup and hips. The blanket normally occurs over, but is not limited to, the hip area.
77:
breeds, though its presence in breeds from Asia to western Europe has indicated that it is due to a very ancient
342:
horses with less than extensive white patterning at birth possess dense white blankets and are called "snowcap."
759:
755:
803:
and the related South German
Coldblood. The existence of leopard-spotted coats among Asian breeds such as the
1447:
Based on these data, we conclude that a susceptibility allele for ERU in
Appaloosas exists in the MHC region.
1975:
628:
was assigned to equine chromosome 1 (ECA1) by a team of researchers. Four years later, this team mapped the
1894:
663:
130:
1727:
1683:
813:
645:
406:
A solid white area normally covering, but not limited to, the hip area, with a contrasting base color.
1267:
742:
horses. A 2008 study theorizes that both CSNB and leopard complex spotting patterns are linked to the
788:
670:
populations. The ancient origin of the allele may explain the presence of spotted horse paintings in
641:
598:
168:
gene unless it is contributed by the other parent. Such horses are termed "non-characteristic" among
57:
spots on a white coat. Secondary characteristics associated with the leopard complex include a white
824:
783:
is the breed best known for the leopard complex patterns, though the complex also characterizes the
1780:
1587:
703:
678:
1884:
1657:
1523:
273:
This horse likely began life completely black, and has since undergone extensive varnish roaning.
208:
110:
666:. This finding represents evidence for the presence of leopard complex spotting in prehistoric
1493:
1485:
1320:
1255:
1148:
1103:
1017:
882:
796:
54:
34:
554:
refers to a horse with a roan blanket which has white and/or dark spots within the roan area.
1879:
1577:
1436:
1310:
1300:
1245:
1235:
1140:
1093:
1085:
987:
874:
758:(ERU) is also present in the breed. Appaloosas have an eightfold greater risk of developing
1534:
1224:"Genotypes of predomestic horses match phenotypes painted in Paleolithic works of cave art"
1012:
Sponenberg, Dan
Phillip (April 11, 2003) . "5/Patterns Characterized by Patches of White".
978:
Sponenberg, D. Phillip (1982). "The inheritance of leopard spotting in the
Noriker horse".
762:(ERU) than all other breeds combined. Up to 25% of all horses with ERU may be Appaloosas.
576:
1506:
991:
792:
780:
425:
169:
106:
50:
421:
Generic term for a horse which has white or dark spots over all or a portion of its body.
41:. These patterns range from progressive increases in interspersed white hair similar to
1722:
1693:
1673:
1632:
1511:
1315:
1288:
1250:
1223:
1098:
1073:
212:
204:
1037:
1999:
1899:
1869:
1854:
1666:
1555:
1441:
1424:
1144:
878:
808:
800:
711:
698:
223:
gene, though a combination of pedigree knowledge and coat characteristics also help.
122:
102:
1402:
1352:
593:
for "leopard complex" by Dr. D. Phillip
Sponenberg in 1982, and was described as an
21:
1930:
1829:
1812:
1802:
1732:
1678:
1547:
784:
614:
504:
495:
466:
A mostly white horse with a bit of color remaining around the flank, neck and head.
291:
282:
278:
98:
94:
74:
46:
42:
455:
164:
gene do not exhibit leopard-complex traits, and cannot produce offspring with the
1305:
1287:
Bellone, Rebecca R.; Holl, Heather; Setaluri, Vijayasaradhi; et al. (2013).
1925:
1889:
1705:
1594:
1518:
1501:
1089:
671:
659:
543:
485:
90:
243:
1874:
1759:
1652:
1617:
1612:
1607:
1564:
841:
829:
750:
694:
682:
667:
640:
or Melastatin 1 (MLSN1). The leopard complex allele contains a 1378 base-pair
606:
188:
62:
1935:
1904:
1808:
1240:
707:
137:
70:
1324:
1259:
1152:
1107:
886:
470:
440:
558:
522:
A fewspot leopard that is completely white with only mottled skin showing.
1970:
1834:
1775:
1637:
1582:
804:
674:
594:
216:
153:
145:
78:
66:
1339:
Electroretinography of congenital night blindness in an Appaloosa filly.
1940:
1864:
1859:
1770:
1739:
1222:
Pruvost, M.; Bellone, R.; Benecke, N.; et al. (November 7, 2011).
1072:
Bellone, Rebecca R; Brooks SA; Sandmeyer L; et al. (August 2008).
763:
715:
613:) were usually "fewspots", while those with a single copy of the gene (
410:
306:
253:
945:
511:
1744:
1642:
1454:
768:
655:
195:
58:
526:
176:
gene possess, at the very least, leopard-complex "characteristics":
710:
does not reach the brain. Rod cells in the retina are connected to
269:
1950:
1717:
823:
743:
637:
575:
305:
268:
242:
38:
25:
One of many patterns for which the leopard complex is responsible.
20:
1268:"Ancient DNA provides new insights into cave paintings of horses"
1965:
1960:
833:
586:
180:
133:
1458:
840:
The approximately 25,000-year-old paintings "Dappled Horses of
734:
has now been localized encodes a protein that channels calcium
735:
873:(2). International Society for Animal Genetics: 134–137.
787:, as well as breeds related to the Appaloosa such as the
605:) were solid-colored, those with two copies of the gene (
129:) controls the presence of leopard-spotting in horses. A
101:. Colored spots reflect the underlying coat color, be it
1341:
Journal of Equine Medicine and Surgery 1977; 1: 226–229.
771:
that makes the breed more susceptible to the disease.
730:
horses were free from the disorder. The gene to which
726:
subjects with CSNB, while all heterozygotes and non-
1913:
1847:
1822:
1793:
1753:
1704:
1691:
1665:
1563:
1554:
1542:
1492:
136:requires only a single copy to produce an affected
1353:"Shedding Light on Night Blindness in Appaloosas"
986:(73). The American Genetic Association: 357–359.
621:) ranged from mere mottled skin to full leopard.
160:(homozygous dominant). Horses without a dominant
1007:
1005:
1003:
1001:
906:
904:
902:
900:
898:
896:
294:, whose breed associations find it undesirable.
1228:Proceedings of the National Academy of Sciences
1203:
1201:
1199:
1197:
1195:
1193:
1191:
1217:
1215:
1213:
1016:(2 ed.). Wiley-Blackwell. pp. 93–4.
1470:
1165:Based on images from Sponenberg 2003, 153-156
1084:(4). Genetics Society of America: 1861–1870.
681:the leopard pattern may have been helpful as
8:
795:. The gene is also relatively common in the
1181:
1179:
1177:
1175:
1173:
1171:
1067:
1065:
1063:
1061:
1059:
1710:
1701:
1569:
1560:
1477:
1463:
1455:
1125:
1123:
1121:
1119:
1117:
380:
1440:
1403:"Uveitis: Medical and Surgical Treatment"
1314:
1304:
1249:
1239:
1097:
940:
938:
973:
971:
969:
967:
580:Probable European wild horse coat colors
1378:. The Appaloosa Project. Archived from
852:
648:which disrupts transcription of TRMP1.
226:While both heterozygous and homozygous
860:
858:
856:
589:was responsible. This gene was termed
172:aficionados. Horses with at least one
992:10.1093/oxfordjournals.jhered.a109669
828:Replica of horses cave painting from
7:
634:transient receptor potential channel
1038:"Guide to Identifying an Appaloosa"
1374:Sandmeyer, Lynne (July 28, 2008).
33:is a group of genetically related
14:
911:Sheila Archer (August 31, 2008).
1533:
1442:10.1111/j.1463-5224.2005.00442.x
1401:Loving, Nancy (April 19, 2008).
1376:"Equine Recurrent Uveitis (ERU)"
1145:10.1111/j.1463-5224.2007.00572.x
879:10.1111/j.1365-2052.2004.01113.x
677:. It is thought that during the
651:In 2011, a study identified the
557:
542:
525:
510:
484:
469:
454:
439:
424:
409:
1351:Oke, Stacey (August 31, 2008).
1337:Witzel CA, Joyce JR, Smith EL.
1185:Identifying the Appaloosa Horse
685:against the snowy environment.
601:gene. Horses without the gene (
382:
1:
1435:(6): 437–450. November 2005.
211:, and freckled skin with the
1306:10.1371/journal.pone.0078280
913:"Studies Currently Underway"
298:Interactions and terminology
1090:10.1534/genetics.108.088807
2022:
1987:Category:Horse coat colors
1956:Endothelin receptor type B
1921:Equine coat color genetics
1014:Equine Coat Color Genetics
748:
265:Leopard-associated roaning
207:, striped hooves with the
1984:
1713:
1603:
1572:
1531:
1429:Veterinary Ophthalmology
1133:Veterinary Ophthalmology
760:Equine Recurrent Uveitis
756:Equine Recurrent Uveitis
235:smaller, scarcer spots.
85:Leopard complex patterns
61:around the eye, striped
1976:Melanocortin 1 receptor
1241:10.1073/pnas.1108982108
980:The Journal of Heredity
699:ophthalmologic disorder
551:Roan Blanket With Spots
1207:Sponenberg 2003, p. 90
1044:. Appaloosa Horse Club
837:
693:Congenital stationary
581:
361:horses and homozygous
311:
274:
248:
26:
16:Coat pattern in horses
1914:Genetics and breeding
1755:Base color variations
1684:Lethal white syndrome
917:The Appaloosa Project
827:
814:foundation bloodstock
579:
309:
272:
246:
219:can now identify the
24:
816:for a given breed.
789:Pony of the Americas
642:long terminal repeat
1274:. November 7, 2011.
1234:(46): 18626–18630.
704:electroretinography
599:incomplete dominant
205:white face markings
123:incomplete dominant
1885:Primitive markings
1486:Equine coat colors
923:on August 24, 2008
838:
664:prehistoric horses
582:
433:Blanket with spots
312:
275:
249:
209:Silver dapple gene
27:
2006:Horse coat colors
1993:
1992:
1843:
1842:
1789:
1788:
1629:
1628:
1625:
1624:
1605:double dilution:
1574:single dilution:
1382:on April 26, 2009
1023:978-0-8138-0759-1
832:displayed in the
565:
564:
91:spotting patterns
2013:
1880:Point coloration
1823:Roaning patterns
1763:
1711:
1702:
1649:
1591:
1570:
1561:
1537:
1526:
1515:
1494:Base coat colors
1479:
1472:
1465:
1456:
1450:
1449:
1444:
1421:
1415:
1414:
1412:
1410:
1398:
1392:
1391:
1389:
1387:
1371:
1365:
1364:
1362:
1360:
1348:
1342:
1335:
1329:
1328:
1318:
1308:
1284:
1278:
1275:
1263:
1253:
1243:
1219:
1208:
1205:
1186:
1183:
1166:
1163:
1157:
1156:
1127:
1112:
1111:
1101:
1069:
1054:
1053:
1051:
1049:
1034:
1028:
1027:
1009:
996:
995:
975:
962:
961:
959:
957:
942:
933:
932:
930:
928:
919:. Archived from
908:
891:
890:
862:
561:
546:
529:
514:
488:
473:
463:Few Spot Leopard
458:
443:
428:
413:
381:
239:White patterning
117:Leopard spotting
49:to distinctive,
2021:
2020:
2016:
2015:
2014:
2012:
2011:
2010:
1996:
1995:
1994:
1989:
1980:
1909:
1839:
1818:
1805:
1795:Leopard complex
1785:
1764:
1757:
1756:
1749:
1698:
1696:
1687:
1661:
1640:
1621:
1599:
1586:(also known as
1585:
1550:
1538:
1529:
1521:
1510:(also known as
1509:
1488:
1483:
1453:
1423:
1422:
1418:
1408:
1406:
1400:
1399:
1395:
1385:
1383:
1373:
1372:
1368:
1358:
1356:
1350:
1349:
1345:
1336:
1332:
1286:
1285:
1281:
1266:
1221:
1220:
1211:
1206:
1189:
1184:
1169:
1164:
1160:
1139:(10): 368–375.
1129:
1128:
1115:
1071:
1070:
1057:
1047:
1045:
1036:
1035:
1031:
1024:
1011:
1010:
999:
977:
976:
965:
955:
953:
952:. December 2002
944:
943:
936:
926:
924:
910:
909:
894:
867:Animal Genetics
864:
863:
854:
850:
822:
793:Colorado Ranger
781:Appaloosa horse
777:
753:
695:night blindness
691:
662:collected from
574:
535:
498:
494:
493:Appaloosa Roan,
402:
376:
349:and homozygous
300:
267:
241:
170:Appaloosa horse
119:
87:
31:leopard complex
17:
12:
11:
5:
2019:
2017:
2009:
2008:
1998:
1997:
1991:
1990:
1985:
1982:
1981:
1979:
1978:
1973:
1968:
1963:
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1953:
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1943:
1938:
1933:
1928:
1923:
1917:
1915:
1911:
1910:
1908:
1907:
1902:
1897:
1892:
1887:
1882:
1877:
1872:
1867:
1862:
1857:
1851:
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1844:
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1838:
1837:
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1783:
1778:
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1765:
1754:
1751:
1750:
1748:
1747:
1742:
1737:
1736:
1735:
1730:
1725:
1723:Splashed white
1714:
1708:
1706:Pinto patterns
1699:
1694:Horse markings
1692:
1689:
1688:
1686:
1681:
1676:
1674:Dominant white
1671:
1669:
1663:
1662:
1660:
1655:
1650:
1635:
1630:
1627:
1626:
1623:
1622:
1620:
1615:
1610:
1604:
1601:
1600:
1598:
1597:
1592:
1580:
1573:
1567:
1558:
1556:Dilution genes
1552:
1551:
1546:
1544:
1540:
1539:
1532:
1530:
1528:
1527:
1516:
1504:
1498:
1496:
1490:
1489:
1484:
1482:
1481:
1474:
1467:
1459:
1452:
1451:
1416:
1393:
1366:
1343:
1330:
1299:(10): e78280.
1279:
1277:
1276:
1209:
1187:
1167:
1158:
1113:
1055:
1029:
1022:
997:
963:
934:
892:
851:
849:
846:
821:
818:
807:and Mongolian
776:
773:
690:
687:
646:retroviral DNA
573:
566:
563:
562:
555:
552:
548:
547:
540:
537:
531:
530:
523:
520:
516:
515:
508:
500:
490:
489:
482:
479:
475:
474:
467:
464:
460:
459:
452:
449:
445:
444:
437:
434:
430:
429:
422:
419:
415:
414:
407:
404:
398:
397:
392:
387:
375:
372:
371:
370:
355:
343:
336:
328:
321:
299:
296:
266:
263:
240:
237:
213:Champagne gene
200:
199:
192:
185:
118:
115:
86:
83:
15:
13:
10:
9:
6:
4:
3:
2:
2018:
2007:
2004:
2003:
2001:
1988:
1983:
1977:
1974:
1972:
1969:
1967:
1964:
1962:
1959:
1957:
1954:
1952:
1949:
1947:
1944:
1942:
1939:
1937:
1934:
1932:
1929:
1927:
1924:
1922:
1919:
1918:
1916:
1912:
1906:
1903:
1901:
1898:
1896:
1893:
1891:
1888:
1886:
1883:
1881:
1878:
1876:
1873:
1871:
1868:
1866:
1863:
1861:
1858:
1856:
1855:Bend-Or spots
1853:
1852:
1850:
1846:
1836:
1833:
1831:
1828:
1827:
1825:
1821:
1815:
1814:
1810:
1804:
1801:
1800:
1798:
1796:
1792:
1782:
1779:
1777:
1774:
1772:
1769:
1768:
1766:
1761:
1752:
1746:
1743:
1741:
1738:
1734:
1731:
1729:
1726:
1724:
1721:
1720:
1719:
1716:
1715:
1712:
1709:
1707:
1703:
1700:
1695:
1690:
1685:
1682:
1680:
1677:
1675:
1672:
1670:
1668:
1664:
1659:
1658:Silver dapple
1656:
1654:
1651:
1647:
1646:
1639:
1636:
1634:
1631:
1619:
1616:
1614:
1611:
1609:
1606:
1602:
1596:
1593:
1589:
1584:
1581:
1579:
1576:
1575:
1571:
1568:
1566:
1562:
1559:
1557:
1553:
1549:
1545:
1541:
1536:
1525:
1520:
1517:
1513:
1508:
1505:
1503:
1500:
1499:
1497:
1495:
1491:
1487:
1480:
1475:
1473:
1468:
1466:
1461:
1460:
1457:
1448:
1443:
1438:
1434:
1430:
1426:
1420:
1417:
1404:
1397:
1394:
1381:
1377:
1370:
1367:
1354:
1347:
1344:
1340:
1334:
1331:
1326:
1322:
1317:
1312:
1307:
1302:
1298:
1294:
1290:
1283:
1280:
1273:
1269:
1265:
1264:
1261:
1257:
1252:
1247:
1242:
1237:
1233:
1229:
1225:
1218:
1216:
1214:
1210:
1204:
1202:
1200:
1198:
1196:
1194:
1192:
1188:
1182:
1180:
1178:
1176:
1174:
1172:
1168:
1162:
1159:
1154:
1150:
1146:
1142:
1138:
1134:
1126:
1124:
1122:
1120:
1118:
1114:
1109:
1105:
1100:
1095:
1091:
1087:
1083:
1079:
1075:
1068:
1066:
1064:
1062:
1060:
1056:
1043:
1039:
1033:
1030:
1025:
1019:
1015:
1008:
1006:
1004:
1002:
998:
993:
989:
985:
981:
974:
972:
970:
968:
964:
951:
947:
941:
939:
935:
922:
918:
914:
907:
905:
903:
901:
899:
897:
893:
888:
884:
880:
876:
872:
868:
861:
859:
857:
853:
847:
845:
843:
835:
831:
826:
819:
817:
815:
810:
806:
802:
798:
794:
790:
786:
782:
774:
772:
770:
765:
761:
757:
752:
747:
745:
741:
737:
733:
729:
725:
719:
717:
713:
712:bipolar cells
709:
705:
700:
696:
689:Vision issues
688:
686:
684:
680:
676:
673:
669:
665:
661:
657:
654:
649:
647:
644:insertion of
643:
639:
635:
631:
627:
622:
620:
616:
612:
608:
604:
600:
596:
592:
588:
578:
571:
567:
560:
556:
553:
550:
549:
545:
541:
538:
533:
532:
528:
524:
521:
518:
517:
513:
509:
506:
501:
497:
492:
491:
487:
483:
480:
477:
476:
472:
468:
465:
462:
461:
457:
453:
450:
447:
446:
442:
438:
435:
432:
431:
427:
423:
420:
417:
416:
412:
408:
405:
400:
399:
396:
393:
391:
388:
386:
383:
379:
373:
368:
364:
360:
357:Heterozygous
356:
352:
348:
345:Heterozygous
344:
341:
337:
333:
329:
326:
323:Heterozygous
322:
318:
315:Heterozygous
314:
313:
308:
304:
297:
295:
293:
292:Knabstruppers
288:
284:
280:
271:
264:
262:
260:
255:
245:
238:
236:
233:
229:
224:
222:
218:
214:
210:
206:
197:
193:
190:
186:
182:
179:
178:
177:
175:
171:
167:
163:
159:
155:
151:
147:
143:
139:
135:
132:
128:
124:
116:
114:
112:
108:
104:
100:
96:
92:
84:
82:
80:
76:
72:
68:
64:
60:
56:
52:
48:
44:
40:
36:
35:coat patterns
32:
23:
19:
1971:STX17 (Gray)
1931:Horse genome
1813:Knabstrupper
1806:
1803:Varnish roan
1794:
1697:and patterns
1679:Sabino-white
1644:
1643:Blue dun or
1446:
1432:
1428:
1419:
1407:. Retrieved
1396:
1384:. Retrieved
1380:the original
1369:
1357:. Retrieved
1346:
1338:
1333:
1296:
1292:
1282:
1271:
1231:
1227:
1161:
1136:
1132:
1081:
1077:
1048:November 12,
1046:. Retrieved
1042:Registration
1041:
1032:
1013:
983:
979:
954:. Retrieved
950:Equine Color
949:
925:. Retrieved
921:the original
916:
870:
866:
839:
785:Knabstrupper
778:
754:
739:
731:
727:
723:
720:
692:
652:
650:
629:
625:
623:
618:
615:heterozygous
610:
602:
590:
583:
569:
534:Roan Blanket
496:Varnish roan
394:
389:
384:
377:
366:
362:
358:
350:
346:
339:
331:
324:
316:
301:
286:
276:
258:
250:
231:
227:
225:
220:
201:
173:
165:
161:
157:
154:heterozygous
149:
148:recessive),
141:
126:
120:
88:
75:Knabstrupper
30:
28:
18:
1926:Color breed
1781:Tricoloured
1758:(primarily
1618:Smoky cream
1595:Smoky black
1405:. The Horse
1359:February 7,
1355:. The Horse
946:"Appaloosa"
927:November 4,
820:In cave art
672:paleolithic
660:DNA samples
390:Description
338:Homozygous
335:(leopard)."
330:Homozygous
279:graying out
109:, gray, or
1760:UK English
1641:(see also
1588:Isabelline
1524:Seal brown
956:October 4,
848:References
842:Pech Merle
830:Pech Merle
775:Prevalence
751:Nyctalopia
749:See also:
683:camouflage
668:wild horse
632:gene to a
607:homozygous
401:Blanket or
146:homozygous
121:A single,
1905:Tiger eye
1809:Appaloosa
1807:See also
1633:Champagne
1522:included
1409:March 21,
1386:March 21,
797:Falabella
708:rod cells
624:In 2004,
595:autosomal
499:or Marble
478:Snowflake
283:true roan
138:phenotype
99:true roan
71:Appaloosa
51:Dalmatian
2000:Category
1951:KIT gene
1936:Wildtype
1900:Mushroom
1835:Rabicano
1776:Skewbald
1608:Cremello
1583:Palomino
1578:Buckskin
1507:Chestnut
1325:24167615
1293:PLOS ONE
1272:Phys.org
1260:22065780
1153:17970998
1108:18660533
1078:Genetics
887:15025575
805:Karabair
675:cave art
536:or Frost
374:Patterns
320:leopard.
217:DNA test
187:striped
184:mottled,
131:dominant
107:chestnut
79:mutation
1941:Melanin
1875:Pangaré
1865:Cropout
1860:Brindle
1771:Piebald
1740:Tobiano
1613:Perlino
1316:3805535
1251:3219153
1099:2516064
801:Noriker
764:Uveitis
718:(Ca).
716:calcium
679:Ice Age
519:Mottled
448:Leopard
403:snowcap
385:Pattern
254:tobiano
95:graying
67:mottled
55:leopard
43:graying
1946:Agouti
1895:Flaxen
1745:Tovero
1733:Sabino
1645:Grullo
1512:Sorrel
1323:
1313:
1258:
1248:
1151:
1106:
1096:
1020:
885:
836:museum
799:, the
769:allele
746:gene.
697:is an
656:allele
636:gene,
259:PATN-1
257:gene,
196:sclera
194:white
189:hooves
156:), or
125:gene (
111:silver
63:hooves
59:sclera
53:-like
39:horses
1890:Sooty
1870:Liver
1848:Other
1728:Frame
1718:Overo
1667:White
1653:Pearl
1565:Cream
1502:Black
809:Altai
744:TRPM1
740:Lp/Lp
638:TRPM1
418:Spots
395:Image
367:Lp/Lp
363:Lp/Lp
359:Lp/lp
354:grow.
351:Lp/Lp
347:Lp/lp
340:Lp/Lp
332:Lp/Lp
325:Lp/lp
317:Lp/lp
158:Lp/Lp
150:Lp/lp
142:lp/lp
103:black
1966:PAX3
1961:MITF
1830:Roan
1811:and
1548:Gray
1543:Gray
1411:2010
1388:2010
1361:2009
1321:PMID
1256:PMID
1149:PMID
1104:PMID
1050:2008
1018:ISBN
958:2008
929:2008
883:PMID
834:Brno
791:and
779:The
736:ions
619:Lplp
611:LpLp
603:lplp
587:gene
572:gene
568:The
505:roan
215:. A
181:skin
134:gene
73:and
65:and
47:roan
29:The
1638:Dun
1519:Bay
1437:doi
1311:PMC
1301:doi
1246:PMC
1236:doi
1232:108
1141:doi
1094:PMC
1086:doi
1082:179
988:doi
875:doi
658:in
617:or
609:or
281:or
97:or
45:or
37:in
2002::
1445:.
1431:.
1427:.
1319:.
1309:.
1295:.
1291:.
1270:.
1254:.
1244:.
1230:.
1226:.
1212:^
1190:^
1170:^
1147:.
1135:.
1116:^
1102:.
1092:.
1080:.
1076:.
1058:^
1040:.
1000:^
982:.
966:^
948:.
937:^
915:.
895:^
881:.
871:35
869:.
855:^
732:Lp
728:Lp
724:Lp
653:Lp
630:Lp
626:Lp
597:,
591:Lp
570:Lp
287:Lp
232:Lp
228:Lp
221:Lp
174:Lp
166:Lp
162:Lp
127:Lp
105:,
81:.
1762:)
1648:)
1590:)
1514:)
1478:e
1471:t
1464:v
1439::
1433:8
1413:.
1390:.
1363:.
1327:.
1303::
1297:8
1262:.
1238::
1155:.
1143::
1137:6
1110:.
1088::
1052:.
1026:.
994:.
990::
984:5
960:.
931:.
889:.
877::
507:.
198:.
191:,
152:(
144:(
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.