Global Genetic Diversity, Introgression, and Evolutionary Adaptation of Indicine Cattle Revealed by Whole Genome Sequencing
Open accessAbstract
Indicine cattle, also referred to as zebu (Bos taurus indicus), play a central role in pastoral communities across a wide range of agro-ecosystems, from extremely hot semiarid regions to hot humid tropical regions. However, their adaptive genetic changes following their dispersal into East Asia from the Indian subcontinent have remained poorly documented. Here, we characterize their global genetic diversity using high-quality whole-genome sequencing data from 354 indicine cattle of 57 breeds/populations, including major indicine phylogeographic groups worldwide. We reveal their probable migration into East Asia was along a coastal route rather than inland routes and we detected introgression from other bovine species. Genomic regions carrying morphology-, immune-, and heat-tolerance-related genes underwent divergent selection according to Asian agro-ecologies. We identify distinct sets of loci that contain promising candidate variants for adaptation to hot semi-arid and hot humid tropical ecosystems. Our results indicate that the rapid and successful adaptation of East Asian indicine cattle to hot humid environments was promoted by localized introgression from banteng and/or gaur. Our findings provide insights into the history and environmental adaptation of indicine cattle.
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The molecular divergence estimate strongly supports two independent domestication events involving distinct aurochs subspecies.
“strongly supporting the two independent domestication events involving distinct aurochs subspecies.”
From The Genomic Landscape of Cattle: Domestication, Dispersal, and Adaptive Evolution (2026) · cites this paper as [10]
The Results section, “Demographic history of indicine cattle” (page unavailable), estimates that indicine groups diverged from taurine groups 251.5–301.2 thousand years ago. The Introduction describes separate taurine and indicine domestications, but identifies the resulting cattle—not their aurochs ancestors—as distinct subspecies. It does not explicitly conclude that the divergence estimate strongly supports domestication of distinct aurochs subspecies. (LLM, conf=0.85)
Experimental — accuracy not yet measuredMolecular divergence of taurine and indicine lineages occurred approximately 201,000-301,200 years ago.
“Molecular divergence of taurine and indicine (also referred to as zebu) lineages is estimated to have occurred at approximately 201,000–301,200 years ago [5,7,8,9,10], strongly supporting the two independent domestication events involving distinct aurochs subspecies.”
From The Genomic Landscape of Cattle: Domestication, Dispersal, and Adaptive Evolution (2026) · cites this paper as [10]
The Results section, “Demographic history of indicine cattle” (page not available), estimates that indicine groups diverged from taurine groups 251,500–301,200 years ago. This supports the divergence claim and its upper bound, but not the cited lower bound of 201,000 years. (LLM, conf=0.90)
Experimental — accuracy not yet measuredThe divergence of Y3a from Y3b, as well as sub-haplogroups within indicine mitogenomes, occurred approximately 23,100-24,800 years ago, while South Asian indicine ancestry diverged from East Asian and African indicine ancestry approximately 10,300-40,100 and 11,800 years ago, respectively.
“The divergence of Y3a from Y3b, as well as sub-haplogroups within indicine mitogenomes, occurred approximately 23,100–24,800 years ago, while South Asian indicine ancestry diverged from East Asian and African indicine ancestry approximately 10,300–40,100 and 11,800 years ago, respectively [9,10].”
From The Genomic Landscape of Cattle: Domestication, Dispersal, and Adaptive Evolution (2026) · cites this paper as [10]
Split into 2 atoms — 0 confirmed, 2 partially supported, 0 disputed, 0 misattributed, 0 unverifiable. Expand the claim to see per-atom verdicts.
Experimental — accuracy not yet measuredThe introgression proportions of banteng, gaur, and yak ancestries into East Asian indicine or Tibetan taurine cattle ranged from 1.13% to 10.21%, 2.06% to 9.98%, and 0.05% to 2.94%, respectively.
“The introgression proportions of banteng (Bos javanicus), gaur (Bos gaurus), and yak (Bos mutus) ancestries into East Asian indicine or Tibetan taurine cattle ranged from 1.13% to 10.21%, 2.06% to 9.98%, and 0.05% to 2.94%, respectively [9,10].”
From The Genomic Landscape of Cattle: Domestication, Dispersal, and Adaptive Evolution (2026) · cites this paper as [10]
The Results section, “The indicine adaptation to the tropical, humid Asian environment” (page unavailable), gives the stated banteng and gaur ancestry ranges for East Asian indicine genomes. The copy includes yak genomes in the analysis, but the searched passages do not report the claimed 0.05%–2.94% yak ancestry range in Tibetan taurine cattle. (LLM, conf=0.88)
Experimental — accuracy not yet measuredA previous study reported an expansion of cattle carrying the I1a sub-haplogroup around 3730 years ago, indicating a marked increase in zebu cattle dispersal and exchange frequency beginning approximately 4000 years ago.
“A previous study also reported an expansion of cattle carrying I1a sub-haplogroup around 3730 years ago [10], indicating a marked increase in zebu cattle dispersal and exchange frequency beginning approximately 4000 years ago.”
From The Genomic Landscape of Cattle: Domestication, Dispersal, and Adaptive Evolution (2026) · cites this paper as [10]
The Results (Demographic history of indicine cattle; page unavailable) report an I1a population increase at approximately 3.73 kya. The Discussion links I1a expansion to indicine entry into East Asia around 3–5 kya. The paper does not establish a marked increase in the frequency of cattle dispersal and exchange beginning around 4 kya. (LLM, conf=0.87)
Experimental — accuracy not yet measured
Indochinese and Chinese cattle populations are dominant in the zebu ancestry, specifically Y-chromosomal sub-haplogroup Y3a and the maternal sub-haplogroup I1a.
“Indochinese and Chinese cattle populations are dominant in the zebu ancestry, specifically Y-chromosomal sub-haplogroup Y3a and the maternal sub-haplogroup I1a [10,60,61].”
From The Genomic Landscape of Cattle: Domestication, Dispersal, and Adaptive Evolution (2026) · cites this paper as [10]
The Results section (‘The Southeast Asian coast was the main entry point of indicine cattle into East Asia’) says Y-chromosomal Y3A3 and maternal I1a were found exclusively in Indochinese and Chinese cattle. The claim names Y3A instead of the more specific Y3A3; the paper reports Y3A as predominant in East Asian indicine and North-Central Chinese cattle, not as the paired sub-haplogroup in that finding. It also does not say that Indochinese and Chinese populations dominate zebu ancestry overall. (LLM, conf=0.84)
Experimental — accuracy not yet measuredGenomic signals of recent selection and adaptive introgression from banteng and gaur into East Asian indicine cattle have been identified; these candidate genes are involved in heat-sensing proteins (TRPA1), paracellular water transport and urine concentration regulation (ILDR), Ca2+ homeostasis (CASR), and resistance to tick-borne diseases (HBA, HBA1, HBQ1, HBM).
“Genomic signals of recent selection and adaptive introgression from banteng and gaur into East Asian indicine cattle have been identified. These candidate genes are involved in multiple functions: heat-sensing proteins (TRPA1), paracellular water transport and urine concentration regulation (ILDR), Ca2+ homeostasis (CASR), and resistance to tick-borne diseases (HBA, HBA1, HBQ1, HBM) [10].”
From The Genomic Landscape of Cattle: Domestication, Dispersal, and Adaptive Evolution (2026) · cites this paper as [10]
Split into 2 atoms — 1 confirmed, 1 partially supported, 0 disputed, 0 misattributed, 0 unverifiable. Expand the claim to see per-atom verdicts.
Experimental — accuracy not yet measuredIn light- and dark-coated South Asian indicine cattle breeds, pigmentation-related genes (LEF1, ASIP) and hair development genes (LIPH, FGF22) were under selection.
“In a comparison of light- and dark-coated South Asian indicine cattle breeds, pigmentation-related genes (LEF1, ASIP) were under selection, as well as hair development genes (LIPH, FGF22) [10].”
From The Genomic Landscape of Cattle: Domestication, Dispersal, and Adaptive Evolution (2026) · cites this paper as [10]
The Results section on South Asian indicine cattle reports selective sweeps near LEF1 and ASIP in light-coated breeds compared with dark-coated breeds. Table 1 in the Results identifies LIPH and FGF22 as hair-development genes under selection, but those findings come from a broader indicine-versus-taurine comparison, not the light-versus-dark South Asian breed comparison. (LLM, conf=0.90)
Experimental — accuracy not yet measuredThe autosomal genetic structure of modern cattle is primarily divided into six ancestries: European taurine, East Asian taurine, African taurine, South Asian or Indian indicine, East Asian or Chinese indicine, and African indicine ancestry.
“The autosomal genetic structure of modern cattle is primarily divided into six ancestries (Table 1): European taurine, East Asian taurine, African taurine, South Asian or Indian indicine, East Asian or Chinese indicine, and African indicine ancestry [9,10,31].”
From The Genomic Landscape of Cattle: Domestication, Dispersal, and Adaptive Evolution (2026) · cites this paper as [10]
Studies based on Y-chromosome microsatellites or SNP markers classify modern cattle paternal origins into taurine Y1, Y2 (Y2a and Y2b), and indicine Y3 (Y3a and Y3b) haplogroups.
“Studies based on Y-chromosome microsatellites or SNP markers classify modern cattle paternal origins into taurine Y1, Y2 (Y2a and Y2b), and indicine Y3 (Y3a and Y3b) haplogroups [9,10].”
From The Genomic Landscape of Cattle: Domestication, Dispersal, and Adaptive Evolution (2026) · cites this paper as [10]
A branch of the Indian zebu I1 haplogroup formed the Chinese zebu I1a sub-haplogroup, while the genome-wide nucleotide diversity was higher in East Asian indicine cattle than in South Asian indicine cattle.
“A branch of the Indian zebu I1 haplogroup formed the Chinese zebu I1a sub-haplogroup, while the genome-wide nucleotide diversity was higher in East Asian indicine cattle than in South Asian indicine cattle [10].”
From The Genomic Landscape of Cattle: Domestication, Dispersal, and Adaptive Evolution (2026) · cites this paper as [10]
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Candidate genes and genomic regions relevant for adaptation to extreme climates in livestock
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Bioinformatics and Computational Biology: Tools for the Genomic Era
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- Authors
- Chen, N., Xia, X., Hanif, Q., Zhang, F., Dang, R., Huang, B., Lyu, Y., Luo, X., Zhang, H., Yan, H., Wang, S., Wang, F., Chen, J., Guan, X., Liu, Y., Li, S., Jin, L., Wang, P., Sun, L., Zhang, J., Liu, J., Qu, K., Cao, Y., Sun, J., Liao, Y., Xiao, Z., Cai, M., Mu, L., Siddiki, A. Z., Asif, M., Mansoor, S., Babar, M. E., Hussain, T., Silva, G. L. L. P., Gorkhali, N. A., Terefe, E., Belay, G., Tijjani, A., Zegeye, T., Gebre, M. G., Ma, Y., Wang, Y., Huang, Y., Lan, X., Chen, H., Migliore, N. R., Colombo, G., Semino, O., Achilli, A., Sinding, M.-H. S., Lenstra, J. A., Cheng, H., Lu, W., Hanotte, O., Han, J., Jiang, Y., & Lei, C.
- Affiliations
- Northwest A&F University (CN)
- National Institute for Biotechnology and Genetic Engineering (PK)
- Yunnan University (CN)
- Chuxiong Normal University (CN)
- Guangxi Vocational University of Agriculture (CN)
- Guangxi Veterinary Research Institute (CN)
- Southwest Forestry University (CN)
- Chattogram Veterinary and Animal Sciences University (BD)
- The University Of Agriculture Dera Ismail Khan (PK)
- Virtual University of Pakistan (PK)
- University of Peradeniya (LK)
- Nepal Agricultural Research Council (NP)
- Arsi University (ET)
- Addis Ababa University (ET)
- Jackson Laboratory (US)
- Haramaya University (ET)
- Ningxia University (CN)
- University of Pavia (IT)
- University of Copenhagen (DK)
- Utrecht University (NL)
- Shandong Academy of Agricultural Sciences (CN)
- Jilin Agricultural University (CN)
- University of Nottingham (GB)
- Chinese Academy of Agricultural Sciences (CN)
- Corresponding author
- Han, Jianlin
- Jiang, Yu
- ORCID
- Chen, Ningbo
- Xia, Xiaoting
- Hanif, Quratulain
- Zhang, Fengwei
- Dang, Ruihua
- Huang, Bizhi
- Lyu, Yang
- Luo, Xiaoyu
- Zhang, Hucai
- Yan, Huixuan
- Wang, Fuwen
- Chen, Jialei
- Guan, Xiwen
- Li, Shuang
- Jin, Liangliang
- Wang, Pengfei
- Sun, Luyang
- Zhang, Jicai
- Liu, Jianyong
- Qu, Kaixing
- Cao, Yanhong
- Sun, Junli
- Liao, Yuying
- Cai, Ming
- Mu, Lan
- Siddiki, Amam Zonaed
- Asif, Muhammad
- Mansoor, Shahid
- Babar, Masroor Ellahi
- Hussain, Tanveer
- Gorkhali, Neena Amatya
- Terefe, Endashaw
- Belay, Gurja
- Tijjani, Abdulfatai
- Zegeye, Tsadkan
- Wang, Yu
- Huang, Yongzhen
- Lan, Xianyong
- Chen, Hong
- Migliore, Nicola Rambaldi
- Colombo, Giulia
- Semino, Ornella
- Achilli, Alessandro
- Sinding, Mikkel-Holger S.
- Lenstra, Johannes A.
- Hanotte, Olivier
- Han, Jianlin
- Jiang, Yu
- Lei, Chuzhao
- Venue
- Nature Communications
- Published
- 2023-11-28
- Volume
- 14
- Issue
- 1
- Pages
- 7803-7803
- ISSN
- 2041-1723 (ISSN-L)
- PMC
- PMC10684552
- OpenAlex
- W4389086892
- Cited by
- 83
- References
- 85
- Citations by year
- 2026: 27 · 2025: 30 · 2024: 26
- FWCI
- 13.95
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Locations
- Nature Communications (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- Repository@Nottingham (University of Nottingham) (publishedVersion, other-oa)
- Research at the University of Copenhagen (University of Copenhagen) (publishedVersion, cc-by)
- RePEc: Research Papers in Economics (submittedVersion)
- CGSPace A Repository of Agricultural Research Outputs (Consultative Group for International Agricultural Research) (submittedVersion, cc-by)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
- Utrecht University Repository (Utrecht University) (submittedVersion, other-oa)
- The Mouseion at the JAXlibrary (Jackson Laboratory) (submittedVersion, cc-by)
- Greater South Information System
- Funders
- Bill and Melinda Gates Foundation (OPP1127286)
- National Natural Science Foundation of China (2021YFD1200400, 202205AR070001, CARS-37, 31872317, 2022YFF1000100, 31861143014, 32072720, 32102523)
- China Postdoctoral Science Foundation
- Chinese Academy of Agricultural Sciences (CARS-37)
- Northwest A and F University (CARS-37)
- Addis Ababa University
- International Livestock Research Institute
- Consortium of International Agricultural Research Centers
- Scotland’s Rural College
- Foreign, Commonwealth and Development Office (OPP1127286)
- Earmarked Fund for Modern Agro-industry Technology Research System (CARS-37)
- Earmarked Fund for China Agriculture Research System (CARS-37)
- National Key Research and Development Program of China (2021YFD1200400, CARS-37, 31861143014, 2022YFF1000100, SQ2021YFF1000041)
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Genetic and phenotypic traits in livestock
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Genetic diversity and population structure
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Genetic Mapping and Diversity in Plants and Animals
- Keywords
- Bos indicus, genetic diversity, introgression, divergent selection, adaptive evolution, heat tolerance, immune-related genes, East Asian migration, phylogeography, banteng, gaur, candidate variants
- MeSH terms
- Whole Genome Sequencing, Alleles, Animals, Cattle, Biological Evolution, Genetic Variation, Ecosystem, Polymorphism, Single Nucleotide
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