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Animals : an open access journal from MDPI2026; 16(2); 297; doi: 10.3390/ani16020297

Differential Expression and Analysis of TBX3 Gene in Skin Tissues of Dun Mongolian Horses with and Without Bider Markings.

Abstract: (1) Background: The dun coat color, a wild-type phenotype in horses characterized by pigment dilution and primitive markings, is regulated by . This study explored the expression and localization of in the Bider marking (a primitive mark unique to the shoulder of horses); (2) Methods: We compared skin tissues from Bider-marked and non-Bider dun Mongolian horses. Samples were collected from the Bider area (dark-colored/light-colored shoulder), dorsal midline, and croup. Histological staining, qRT-PCR, and Western blotting were used to analyze pigment distribution and expression at mRNA and protein levels; (3) Results: Histology revealed asymmetric pigment deposition in hair shafts from light-colored areas of both Bider and non-Bider horses, whereas dark areas showed symmetric distribution. qRT-PCR and Western blotting showed expression was significantly higher in the shoulder of non-Bider horses compared to Bider horses. Conversely, Bider horses exhibited higher levels in all other sampled areas. Immunohistochemistry localized protein to the epidermis and hair follicle bulbs in both groups; (4) Conclusions: In dun Mongolian horses, expression differences between dark and light skin areas correlate with Bider markings. is implicated in this specific pigment marking, though its upstream regulation requires further study. These findings provide key insights into the mechanism behind Bider marking formation.
Publication Date: 2026-01-18 PubMed ID: 41594486PubMed Central: PMC12837385DOI: 10.3390/ani16020297Google Scholar: Lookup
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Summary

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Overview

  • This study investigates the role of the TBX3 gene in the distinct Bider markings found on the skin of dun Mongolian horses, comparing areas with and without these primitive pigment patterns.

Introduction and Background

  • The dun coat color in horses is a natural, wild-type phenotype characterized by a lightened pigment base and primitive markings such as dorsal stripes and leg bars.
  • Bider marking is a unique primitive marking typically found on the shoulder of some dun horses, exhibiting a distinct pattern of pigmentation.
  • TBX3 is a gene associated with pigmentation regulation and was hypothesized to have a role in the formation of these markings.

Research Objectives

  • To analyze differential expression of the TBX3 gene at both mRNA and protein levels in dun Mongolian horses exhibiting Bider markings versus those without.
  • To correlate TBX3 expression with pigment distribution and localize the protein within skin tissue structures.

Methods

  • Skin tissue samples were collected from three distinct areas of the horses:
    • Bider area on the shoulder, distinguished further into dark-colored and light-colored shoulder regions.
    • Dorsal midline area.
    • Croup area (rear part of the horse).
  • Histological staining was used to examine pigment distribution in hair shafts to assess asymmetry or symmetry of pigment deposition.
  • Quantitative real-time PCR (qRT-PCR) was conducted to measure TBX3 mRNA expression levels.
  • Western blotting was performed to quantify TBX3 protein expression in the sampled skin tissues.
  • Immunohistochemistry was employed to localize TBX3 protein within the skin layers and structures such as the epidermis and hair follicle bulbs.

Key Findings

  • Histological analysis showed:
    • Asymmetric pigment deposition in hair shafts from light-colored shoulder areas in both Bider-marked and non-Bider horses.
    • Symmetric pigment distribution in darker-colored areas.
  • Expression patterns of TBX3:
    • Non-Bider horses showed significantly higher TBX3 expression in the shoulder region compared to Bider-marked horses.
    • Bider markings were associated with higher TBX3 expression across other sampled skin areas (dorsal midline and croup) in Bider horses compared to their non-Bider counterparts.
  • The TBX3 protein localized predominantly in:
    • The epidermis, the outer skin layer.
    • The bulbs of hair follicles, which are important sites for pigment-producing melanocytes and hair growth.

Interpretation and Implications

  • The study suggests a direct relationship between TBX3 gene expression levels and the formation of Bider markings in dun Mongolian horses.
  • The variable expression of TBX3 between dark and light skin areas aligns with the visual pigment patterns characteristic of the Bider markings.
  • Localization of TBX3 in key pigment-related skin structures supports its functional role in pigment formation and regulation.
  • These findings demonstrate that TBX3 is likely a critical gene contributing to the specific primitive marking phenotype; however, the upstream mechanisms that regulate TBX3 expression are still unknown and warrant further research.

Conclusions

  • TBX3 plays an important role in differentiating pigment markings on the skin of dun Mongolian horses, particularly relating to the unique Bider phenotype.
  • The study provides foundational insight that can help unravel the genetic and molecular basis of primitive pigmentation patterns in horses.
  • Future directions include identifying the upstream regulatory factors that influence TBX3 expression and understanding how these contribute to complex coat color traits.

Cite This Article

APA
An T, Dugarjaviin M. (2026). Differential Expression and Analysis of TBX3 Gene in Skin Tissues of Dun Mongolian Horses with and Without Bider Markings. Animals (Basel), 16(2), 297. https://doi.org/10.3390/ani16020297

Publication

ISSN: 2076-2615
NlmUniqueID: 101635614
Country: Switzerland
Language: English
Volume: 16
Issue: 2
PII: 297

Researcher Affiliations

An, Tana
  • Inner Mongolia Key Laboratory of Equine Science Research and Technology Innovation, Inner Mongolia Agricultural University, Hohhot 010018, China.
Dugarjaviin, Manglai
  • Inner Mongolia Key Laboratory of Equine Science Research and Technology Innovation, Inner Mongolia Agricultural University, Hohhot 010018, China.

Grant Funding

  • U23A20224 / National Natural Science Joint Fund Key Support Project

Conflict of Interest Statement

The authors declare no conflicts of interest.

References

This article includes 22 references
  1. Gaunitz C, Fages A, Hanghøj K, Albrechtsen A, Khan N, Schubert M, Seguin-Orlando A, Owens IJ, Felkel S, Bignon-Lau O. Ancient genomes revisit the ancestry of domestic and Przewalski’s horses. Science 2018;360:111–114.
    doi: 10.1126/science.aao3297pubmed: 29472442google scholar: lookup
  2. Fages A, Hanghøj K, Khan N, Gaunitz C, Seguin-Orlando A, Leonardi M, McCrory Constantz C, Gamba C, Al-Rasheid KAS, Albizuri S. Tracking Five Millennia of Horse Management with Extensive Ancient Genome Time Series. Cell 2019;177:1419–1435.e31.
    doi: 10.1016/j.cell.2019.03.049pmc: PMC6547883pubmed: 31056281google scholar: lookup
  3. Guimaraes S, Arbuckle BS, Peters J, Adcock SE, Buitenhuis H, Chazin H, Manaseryan N, Uerpmann H-P, Grange T, Geigl E-M. Ancient DNA shows domestic horses were introduced in the southern Caucasus and Anatolia during the Bronze Age. Sci Adv 2020;6:eabb0030.
    doi: 10.1126/sciadv.abb0030pmc: PMC7494339pubmed: 32938680google scholar: lookup
  4. Librado P, Khan N, Fages A, Kusliy MA, Suchan T, Tonasso-Calvière L, Schiavinato S, Alioglu D, Fromentier A, Perdereau A. The origins and spread of domestic horses from the Western Eurasian steppes. Nature 2021;598:634–640.
    doi: 10.1038/s41586-021-04018-9pmc: PMC8550961pubmed: 34671162google scholar: lookup
  5. Librado P, Tressières G, Chauvey L, Fages A, Khan N, Schiavinato S, Calvière-Tonasso L, Kusliy MA, Gaunitz C, Liu X. Widespread horse-based mobility arose around 2200 BCE in Eurasia. Nature 2024;631:819–825.
    doi: 10.1038/s41586-024-07597-5pmc: PMC11269178pubmed: 38843826google scholar: lookup
  6. Ludwig A, Pruvost M, Reissmann M, Benecke N, Brockmann GA, Castaños P, Cieslak M, Lippold S, Llorente L, Malaspinas A-S. Coat color variation at the beginning of horse domestication. Science 2009;324:485.
    doi: 10.1126/science.1172750pmc: PMC5102060pubmed: 19390039google scholar: lookup
  7. Sponenberg DP, Bellone R. Equine Color Genetics. 4th ed. Wiley-Blackwell; Hoboken, NJ, USA: 2017.
  8. Imsland F, McGowan K, Rubin C-J, Henegar C, Sundström E, Berglund J, Schwochow D, Gustafson U, Imsland P, Lindblad-Toh K. Regulatory mutations in TBX3 disrupt asymmetric hair pigmentation that underlies Duncamouflage color in horses. Nat Genet 2016;48:152–158.
    doi: 10.1038/ng.3475pmc: PMC4731265pubmed: 26691985google scholar: lookup
  9. An T, Dugarjaviin M, Han H. Expression and Analysis of TBX3 Gene in the Skin from Three Locations on Dun Mongolian Bider Horse. Genes 2024;15:1589.
    doi: 10.3390/genes15121589pmc: PMC11675668pubmed: 39766856google scholar: lookup
  10. Cieslak J, Brooks SA, Wodas L, Mantaj W, Borowska A, Sliwowska JH, Ziarniak K, Mackowski M. Genetic Background of the Polish Primitive Horse (Konik) Coat Color Variation-New Insight into Dun Dilution Phenotypic Effect. J Hered 2021;112:436–442.
    doi: 10.1093/jhered/esab034pubmed: 34432873google scholar: lookup
  11. Stachurska AM. Inheritance of primitive markings in horses. J Anim Breed Genet 1999;116:29–38.
  12. Masuda M, Tsunoda J, Nomura H, Kimura N, Altangerel G, Namkhai B, Dolj U, Yokohama M. New Primitive Marking (Bider) in Mongolian Native Horse and Equus przewalskii. J Equine Sci 2007;18:145–151.
    doi: 10.1294/jes.18.145google scholar: lookup
  13. Thiruvenkadan AK, Kandasamy N, Panneerselvam S. Coat colour inheritance in horses. Livest Sci 2008;117:109–129.
  14. Khaudov AD, Zhekamukhov MH, Zaitsev AM, Amshokova ZH, Berbekova NV, Amshokov HK. Analysis of ASIP and MC1R gene polymorphisms in Kabardin horses. Russ Agric Sci 2024;50:185–189.
    doi: 10.3103/S1068367424700010google scholar: lookup
  15. Liu X, Peng Y, Zhang X, Wang X, Chen W, Kou X, Liang H, Ren W, Khan MZ, Wang C. Coloration in Equine: Overview of Candidate Genes Associated with Coat Color Phenotypes. Animals 2024;14:1802.
    doi: 10.3390/ani14121802pmc: PMC11200706pubmed: 38929421google scholar: lookup
  16. Lusis JA. Striping markings in domestic horses.. Genetica 1943;23:31–62.
    doi: 10.1007/BF01763802google scholar: lookup
  17. Han H, Bryan K, Shiraigol W, Bai D, Zhao Y, Bao W, Yang S, Zhang W, E MacHugh D, Dugarjaviin M. Refinement of Global Domestic Horse Biogeography Using Historic Landrace Chinese Mongolian Populations.. J. Hered. 2019;110:769–781.
    doi: 10.1093/jhered/esz032pubmed: 31628847google scholar: lookup
  18. Zhang B, Chen T. Local and systemic mechanisms that control the hair follicle stem cell niche.. Nat. Rev. Mol. Cell Biol. 2024;25:87–100.
    doi: 10.1038/s41580-023-00662-3pubmed: 37903969google scholar: lookup
  19. Shang S, Yu Y, Zhao Y, Dang W, Zhang J, Qin X, Irwin DM, Wang Q, Liu F, Wang Z. Synergy between MC1R and ASIP for coat color in horses (Equus caballus). J. Anim. Sci. 2019;97:1578–1585.
    doi: 10.1093/jas/skz071pmc: PMC6447268pubmed: 30785190google scholar: lookup
  20. Ali SA, Naaz I. Biochemical aspects of mammalian melanocytes and the emerging role of melanocyte stem cells in dermatological therapies.. Int. J. Health Sci. 2018;12:69–76.
    pmc: PMC5870308pubmed: 29623021
  21. Moreiras H, Seabra MC, Barral DC. Melanin Transfer in the Epidermis: The Pursuit of Skin Pigmentation Control Mechanisms.. Int. J. Mol. Sci. 2021;22:4466.
    doi: 10.3390/ijms22094466pmc: PMC8123122pubmed: 33923362google scholar: lookup
  22. Shi X, Zhuang Y, Chen Z, Xu M, Kuang J, Sun X-L, Gao L, Kuang X, Zhang H, Li W. Hierarchical deployment of Tbx3 dictates the identity of hypothalamic KNDy neurons to control puberty onset.. Sci. Adv. 2022;8:eabq2987.
    doi: 10.1126/sciadv.abq2987pmc: PMC9668310pubmed: 36383654google scholar: lookup

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