New Primitive Marking (Bider) in Mongolian Native Horse and Equus Przewalskii
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—ORIGINAL— New Primitive Marking (Bider) in Mongolian Native Horse and Equus przewalskii Mioko MASUDA1, Junko TSUNODA2, Hiroko NOMURA2, Nami KIMURA2, Gombojav ALTANGEREL1, Bandi NAMKHAI3, Usukhjargal DOLJ3 and Michinari YOKOHAMA2* 1Mongolian State University of Agriculture, Zaisan-53, Ulaanbaatar-210153, Mongolia,2Faculty of Bioindustry, Tokyo University of Agriculture, Yasaka-196, Abashiri-shi, 099-2493 Hokkaido, Japan, 3Hustai National Park, P.O. Box 1160, Central Post Office, Ulaanbaatar, 13, Mongolia The incidence of Bider (Bi marking) in Mongolian native horses was 0.010, and that J. Equine Sci. according to districts was the highest (0.032) in Tov Aimag. Bi marking was frequently Vol. 18, No. 4 observed in bay (0.482) and chestnut horses (0.351), and was also observed in gray and roan pp. 145–151, 2007 horses. The incidence of Bi marking was high (0.842) in horses with a dorsal strip (DS) as a primitive marking. In 164 Equus przewalskii, the incidence of Bi marking was 0.396, which was about 40 times the incidence in Mongolian native horses. Bi marking was observed in both males and females in the two species, showing autosomal inheritance. Analysis of the mode of inheritance based on the incidence of Bi marking in foals according to various crossing combinations showed the complete dominance of Bi marking in both species. Thus, it was concluded that Bi marking occurs in Bi/Bi and Bi/– genotypes. This Bi marking differs from DS, leg stripes (LS) and shoulder stripes (SS), and is considered to be a type of primitive marking peculiarly occurring in Mongolian native horse and Equus przewalskii. The high incidence of Bi marking in Equus przewalskii (0.396), and the results of crossing between Mongolian native horses and Equus przewalskii performed in Mongolia, suggest introgression of the Bi marking gene from Equus przewalskii to Mongolian native horses. Key words: Bider (Bi), coat color gene, Equus przewalskii, primitive markings, Mongolian native horse The coat color of domestic horses shows variation, coat in Thoroughbreds is expressed by the Sabino- sometimes representing a species’ characteristic. At based white gene (S), and its homozygotes have a white present, 13 coat color-associated genes have been coat. identified [2, 6–8, 11], but new findings have also been In the basic coat, such as bay and chestnut in reported recently. Though homozygosity for the roan domestic horses, the loci of extension (E) and Agouti gene (R) as a coat color-associated gene was reported to (A) as coloring genes are closely involved, and the be lethal in the Belgian breed [7], Ueda [12, 13] and expression of coat color is controlled by melanocortin- Yokohama et al. [14, 15] showed the survival of 1 receptors (MC1R) and agouti-signaling protein Hokkaido native horses with R/R type. It was also (ASIP) genes [5, 10]. In these loci, point mutation of C reported that the white lethal gene (W) is involved in (E gene) at codon 901→T (e gene) in MC1R and the white coat, and that the homozygote white gene is deletion mutation (A gene→a gene) of 11 bp in 2174– also lethal [7], but Brooks and Bailey [1], and 2184 bp in the second exon of ASIP have been Yokohama and Ishida [16] confirmed that the white reported. The E locus is epistatic to the A locus, and chestnut color develops irrespective of the genotype of e e This article was accepted August 27, 2007 the A locus in recessive homozygosity (E /E ). On the a a *Corresponding author. other hand, when the A locus is A /A , a black coat e-mail: [email protected] develops in the presence of at least one dominant allele 146 M. MASUDA, J. TSUNODA, H. NOMURA ET AL. The survey items were the basic color, incidences of primitive markings, and incidence of Bi marking in the parent-child relationship in each district. The number of investigated horses was 3,775 during 1997–2003 and 529 in 2006 (total 4,304). A survey of Equus przewalskii was performed in the Hustai National Park located in the suburbs of Ulaanbaatar, the capital of Mongolia, which is one of the three places designated as Equus przewalskii protection areas. Of a total of 194 horses in the park (June, 2006), 164, in which the presence or absence of Fig. 1. Survey areas (eight Aimags) of Mongolian native horses. Bi marking could be determined, were included in this Remarks: Number showed animal number in each area, and horse number with Bider showed in parentheses. study. Bi markings were confirmed by direct : Other Aimags; n=2,700 (2). observation using telescopes and photography. In horse that could not be observed, Bi markings were confirmed on the basis of literal data and photographs stored in the Hustai National Park archives. in both the A and E loci. In Equidae, DS, LS and SS as primitive markings are Investigation of coat colors and markings observed [9]. These are rare characters in improved In Mongolian native horses, four types of primitive domestic horses but are frequently observed in local marking (DS, LS, SS and Bi) were evaluated (Fig. 2). horse breeds and wild horses. These markings have All Equus przewalskii had DS and LS. linear patterns. However, in our survey in Mongolia in 1998, we observed Mongolian native horses with Bi Sex difference and mode of inheritance of Bi marking marking that had not been observed in domestic In 646 Mongolian native horses and 164 Equus horses. Unlike DS, LS and SS, the Bi marking has an przewalskii, the difference in the incidence of Bi irregular coat color/pattern. Subsequently, in 2003, we marking between males and females was analyzed by discovered in the literature the existence of Equus the χ2 test. Then, based on data on 296 Mongolian przewalskii which has the same marking and habitat of native foals and 164 Equus przewalskii foals, the the in Mongolian grassland [4]. Since the Bi marking theoretical incidences of foals with and those without is a very rare character and is also observed in Equus Bi markings were calculated from the probability of przewalskii, we considered that this marking might be a crossing combinations. The difference between the marker with which to clarify the genetic relationship theoretical and observed incidences was analyzed by between domestic and wild horses. Therefore, a the χ2 test, and the mode of inheritance was evaluated. statistical survey was performed to evaluate the incidence of Bi marking and the mode of its inheritance in Equus przewalskii, which are protected Results and proliferate in Mongolia, and Mongolian native horses. Development of Bider (Bi) markings Bi markings vary in size among individual horses, Materials and Methods exhibiting the small punctate type, the medium-sized belt-like type extending to the area around the shoulder, or the large type extending to the neck and Investigated horses costal region (Fig. 3). Unlike linear markings, the Bi Surveys of the basic coat color and Bi marking in marking has an irregular symmetrical pattern and is Mongolian native horses were performed during 1997– composed of skin tissue pigments, as is in tattoos (Fig. 2003 and 2006 (April–September) in eight Aimags, 4). These findings were similar in Mongolian native mainly in Tov, Hentii and Dundgovi Aimags (Fig. 1). horses and Equus przewalskii (Fig. 5). BIDER MARKING IN MONGOLIAN HORSE 147 Fig. 2. Primitive markings in Equus przewalskii. DS: Dorsal stripe, LS: Leg stripe, SS: Shoulder Stripe, Bi: Bider. Phot. 1 cited from Lekler B.(2003)6). Fig. 3. Variation of Bider (Bi ) type in Mongolian native horses. 148 M. MASUDA, J. TSUNODA, H. NOMURA ET AL. Fig. 4. Bider is an irregular symmetrical pattern. Remarks: Depicted horse has a bay coat color. L: Left side, R: Right side. Fig. 5. Variation of Bider (Bi) patterns in Equus przewalskii in the Hustai National Park located in the suburbs of Ulaanbaatar. The incidences of coat colors and primitive markings in Mon- marking was 0.149, but that of Bi markings golian native horses accompanied by DS was 0.842 (Table 1). These In Mongolian native horses with Bi markings, the findings suggest a close association between Bi and DS basic coat color was classified into 10 types (Table 1). markings. Table 1 shows that the Bi marking is not a character developing for specific coat colors. The incidence of Bi The incidence of Bi marking in Mongolian native horses marking was the highest for bay (0.482), followed by according to districts chestnut (0.351). In this study, the incidence of Bi The incidence of Bi marking in the Mongolian native marking not accompanied by any other primitive horses was 0.010, showing it to be a very rare character BIDER MARKING IN MONGOLIAN HORSE 149 Table 1. Frequency of appearance of coat colors and primitive markings in Mongolian native horses Coat colors Bay Brown Chestnut Dark Palomino Buckskin Gray Roan Bay Chestnut Total Primitive chestnut tabiano tabiano markings Bi 0.421 0.176 0.235 0.000 0.118 0.000 0.059 0.000 0.000 0.000 0.149 (7) (3) (4) (0) (2) (0) (1) (0) (0) (0) (17) Bi/LS 0.375 0.000 0.250 0.125 0.000 0.000 0.000 0.125 0.000 0.125 0.070 (3) (0) (2) (1) (0) (0) (0) (1) (0) (1) (8) Bi/DS 0.400 0.000 0.467 0.067 0.000 0.067 0.000 0.000 0.000 0.000 0.132 (6) (0) (7) (1) (0) (1) (0) (0) (0) (0) (15) Bi/LS/DS 0.521 0.00 0.37 0.00 0.00 0.041 0.041 0.00 0.014 0.014 0.640 (38) (0) (27) (0) (0) (3) (3) (0) (1) (1) (73) Bi/LS/SS 1.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.009 (1) (0) (0) (0) (0) (0) (0) (0) (0) (0) (1) Total 0.482 0.026 0.351 0.017 0.017 0.035 0.035 0.009 0.009 0.018 (114) (55) (3) (40) (2) (2) (4) (4) (1) (1) (2) Bi: Bider, DS: Dorsal Stripe, LS: Leg Stripe, SS: Shoulder Stripe.