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White Markings in Horses

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White Markings in Horses 74 Medycyna Wet. 2012, 68 (2) Artyku³ przegl¹dowy Review White markings in horses ANNA STACHURSKA, ANNE PHAFF USSING* Department of Horse Breeding and Use, Faculty of Biology and Animal Breeding, University of Life Sciences, Akademicka 13, 20-950 Lublin, Poland *The Royal Library, P.O. Box 2149, DK-1016 Copenhagen K, Denmark Stachurska A., Phaff Ussing A. White markings in the horse Summary White markings in horses result from the lack of melanocytes in the skin and hairs. The trait is characteristic of most breeds of domestic horses. In the breeding of the Polish Konik horse, which should resemble its wild ancestors, the lack of white markings was accepted as a selection criterion. The heritability of the markings is high. MC1R and ASIP loci, as well as two other putative QTLs are involved in the appearance of white markings. A foal should inherit a number of genes from both parents to make it possible for the markings to appear. The ultimate extent of markings is caused by the genes, as well as by intrauterine factors. The latter consist of accidental and random events which disturb the survival, migration and clonal proliferation of melanoblasts. The recessive e allele from MC1R locus and the dominant A allele from ASIP locus cause a greater extent of the markings, behaving as major genes. The most desired genotype of the Polish Konik horse, aaEEDD, which in inter se matings produces solely blue dun foals, is also the most desired with regard to the low amount of markings. To increase the progress in the Polish Konik breeding with regard to the absence of the markings, marked individuals and identified carriers of e allele should be consistently eliminated from reproduction. Keywords: white markings, inheritance, horse White markings are patches of white hairs growing melanocytes from the epidermis (24). The patches of on unpigmented skin on the horses peripheral areas: depigmentation usually appear around the anus and the head and limbs. Rarely, the markings appear in genital regions, as well as in the face (fig. 1). Vitiligo central areas, which in the majority of cases indicates is observed in many animal species: mostly mammals, that the horse carries a gene producing one of the pied but also in chickens. The studies performed in humans coat color patterns. The unpigmented skin and hairs show that besides genetic effects it may result from an result from a lack of melanocytes (17). The white autoimmune disorder (20). markings are present at the foals birth and persist In many horse breeds, white markings are assumed throughout the life of the horse. Besides the markings, to be flashy, highlighting the limb action which is vitiligo may develop in adult horses due to the loss of particularly important in shows but not desired in Fig. 1. Vitiligo in Felin Ponies. D¹browica near Lublin Medycyna Wet. 2012, 68 (2) 75 dressage. Moreover, the markings enable the recogni- tion of the horse from a distance. Unpigmented hoof horn associated with the markings is equally hard and elastic as the pigmented one (9, 28). However, because of the unjustified opinion of the inferior quality of the unpigmented horn, horses with white markings are unfavorable in many breeds. Occurrence of white markings Archive sources on the Tarpan (Equus caballus gmelini) living in Poland and Lithuania do not men- tion any white markings. It may be supposed that if the old chronicles name such details as the dorsal stripe, white markings would not be passed over, either. Gmelin (1745-1774) (quoted after 21) states that the wild horses are similar to a mouse with regard to the coat color and their legs beneath the middle to the hooves are black. The lack of white markings in the Fig. 2. Percentage of Polish Konik horses without markings registered in I to VIII volumes of the studbook (24) Przewalski horse (Equus caballus przewalskii), the last living wild representative of the species, and in the Paleolithic cave paintings illustrating the horse, as well minary stud book were then included in the main stud as the markings rarely occurring in primeval Asiatic book (26; fig. 2). The number of the horses without horses (12), indicate the trait is characteristic of markings is higher in the following volumes; however, domestication. Hence, in the breeding of the Polish it still only slightly exceeds 80%. These data show that Konik horse, which should resemble its wild ancestors, the constant occurrence of marked horses, despite the the absence of white markings was accepted as a se- selection, is a great problem in the breeding of Polish lection criterion. Similarly, in the Norwegian Fjord Koniks. horse which is bred for a yellow dun color, white mar- kings seldom occur and are undesired (29). A facial Inheritance of white markings star, up to 5 cm diameter, is solely accepted in brood For over 80 years geneticists have been studying mares, but not in stallions, in Norway. The reason for the inheritance of white markings. In the beginning, the official acceptance is that such a marking was pre- the most extensive research performed in Germany sent in three famous Fjord ancestors in the nineteenth enabled the suggestion of the hypotheses of two pairs century. In Denmark, only white hairs on the forehead of genes controlling the trait (18, 30). Crew and in Fjords are accepted. Buchannan Smith (8) put forward a theory of a domi- Breeders of Polish Konik, Fjord and other horses nant M factor and modifiers. Afterwards, German stu- which should be without white markings always face dies assumed four genes responsible for the markings the dilemma if it is worth eliminating an otherwise (3-6). According to another hypothesis the markings valuable horse due to, for instance, a small white spot. were considered as the least developed pied pattern Hence, usually some exceptions are considered. In (2). Following that approach, Lauvergne et al. (15) Polish Konik breeding, two programs are in effect. suggested horse classification related to the extent of According to the Programme of Conservation Breeding white patches on the body. None of the hypotheses were accepted by the Minister of Agriculture in 1999, confirmed later. Polish Koniks ( ) without white markings may be In the early 1980s, monozygotic horse twins produ- registered in the studbook ( ) temporary in mares ced by embryo micromanipulation attracted the gene- little markings on head (white hairs, snip) are allowed. ticists attention (1). One colt had a marking on all The Breeding Programme of Genetic Resources four limbs and another colt had it solely on the left Conservation effective from 2000 is more strict, since limbs. In a pair of fillies one foal had a marking on the it allows only horses without any white markings. left forelimb and the other did not have any markings. Analyzing the stud book, it may be noticed that up to The result was not consistent with expectations, since Volume VI some individuals with a marking, includ- if the trait is genetically controlled in monozygotic ing the fetlock joint, occur. In Volume VII, mares with twins the genotype and consequently the markings a blaze, stripe or white pastern, as well as stallions should be identical (35). A similar riddle was noticed with a star are registered. There are still five stallions in the case of individual horses with asymmetrical with white hairs, one with a snip and two with a star in markings. The same genotype produces differently Volume IX, which is the latest volume. The percentage expressed traits on two sides of the body. The findings of horses without markings is lower in Volumes III indicated that additional non-genetic factors determine and IV due to the fact that individuals from the preli- the extension of the trait. 76 Medycyna Wet. 2012, 68 (2) Extensive studies on the occurrence and amount of markings brought the solution to the problem. A high or medium correlation between the extension of the facial and limb markings (10, 19, 23, 30, 35) showed that they were due to one genetic mechanism. Results documenting a high sire-foal and dam-foal regression proved the mechanism could not be founded on one pair of genes which would cause the segregation in the offspring. It was also found that stallions were slightly more marked than mares, hence a gene producing the trait might be associa- ted with the horses gender (10, 23, 35). A hypothesis of the multifactorial mode of inheritance of white markings was put forward by Nebe (19) on the basis of data collected in over 8,000 foals of Hessian Fig. 3. Vertical black stripes in Polish Konik stallion. The stripes are not saddle horses. Broad investigations of Ara- present on the other side of the body. Roztocze National Park bian horses registered in the American stud- book, conducted by Woolf at the turn of the 1980s and (breed or identity marks) destroys the melanocytes, 1990s (31-36) explored the hypothesis on the multi- resulting in growth of white hairs in the involved area. factorial inheritance of the markings. Particular analy- Factors disturbing melanoblast migration and pro- ses concerned 2.5 thousand to 12 thousand horses. The liferation are not known. Although the effects should markings were scored from 0 to 5 depending on the similarly influence both sides of the organism, stocha- amount of whiteness at five areas on the head and stic events appear. It must be emphasized that the separately on each limb. According to those studies, migration and proliferation of the melanoblasts occur the heritability of the markings on the head is 0.69, on autonomously in each side of the embryo because of the limbs 0.68, and combining the facial score and four the mid-dorsal separation (33).
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