Bulletin of KSAU Bulletin of KSAU Вестник КрасГАУ 1819-4036 106785 10.36718/1819-4036-2026-3-130-142 onatur Зоотехния и ветеринария Animal breeding and veterinary surgery Зоотехния и ветеринария GWAS ANALYSIS OF HOLSTEIN CATTLE IN THE SVERDLOVSK REGION USING FULL AND EXTREME SAMPLES GWAS-АНАЛИЗ ГОЛШТИНСКОГО КРУПНОГО РОГАТОГО СКОТА СВЕРДЛОВСКОЙ ОБЛАСТИ С ИСПОЛЬЗОВАНИЕМ ПОЛНОЙ И ЭКСТРЕМАЛЬНОЙ ВЫБОРОК Богатова Полина Сергеевна Bogatova Polina Sergeevna polina.bogatova.me@gmail.com Лиходеевская Оксана Евгеньевна Lihodeevskaya Oksana Evgenievna lixodeevskaya@mail.ru Лоретц Ольга Геннадьевна Loretts Olga Gennadievna olga-loretts@yandex.ru Лиходеевский Георгий Александрович Lihodeevskiy Georgiy Aleksandrovich georglihodey@gmail.com Минина Ольга Александровна Minina Olga Aleksandrovna olia-al-li@bk.ru Евсеева Татьяна Павловна Evseeva Tatiana Pavlovna tanya_evseeva@bk.ru Военный учебно-научный центр военно-воздушных сил «Военно-воздушная академия имени профессора Н. Е. Жуковского и Ю. А. Гагарина» Россия Military Educational and Scientific Centre of the Air Force «Professor N.E. Zhukovsky and Y.A. Gagarin Air Force Academy» Russian Federation 03 04 2026 03 04 2026 3 130 140 18 11 2025 https://vestnik.kgau.ru/en/nauka/article/106785/view

Цель исследования – выявление генетических маркеров, ассоциированных с показателями молочной продуктивности (удоем, содержанием жира и белка в молоке) у голштинского крупного рогатого скота, с использованием метода полногеномного ассоциативного анализа (GWAS) на выборке животных из племенных хозяйств Свердловской области за 2018–2024 гг. Задачи: провести генотипирование выборки коров с использованием ДНК-биочипов и осуществить контроль качества данных; выполнить GWAS-анализ молочной продуктивности на полной популяции и на экстремальных фенотипических группах; идентифицировать статистически значимые однонуклеотидные полиморфизмы (ОНП), связанные с удоем, содержанием жира и белка; проанализировать гены, расположенные вблизи значимых ОНП, для понимания их роли в метаболизме липидов, иммунном ответе и развитии ткани молочной железы; оценить совпадение генетических ассоциаций между полной выборкой и экстремальными группами для подтверждения надежности маркеров. GWAS-анализ молочной продуктивности проведен на 539 голштинских коровах из 3 племенных хозяйств Свердловской области (2018–2024 гг.). Анализ удоя, жира и белка выполнен на полной выборке и экстремальных квартилях Q1/Q4 (удой: (10028 ± 680) и (6775 ± 653) кг; жир: (4,07 ± 0,068) и (3,79 ± 0,092) %; белок: (3,38 ± 0,066) и (3,12 ± 0,045) %). Всего идентифицировано 20 значимых ОНП, ассоциированных с показателями продуктивности крупного рогатого скота. Вблизи некоторых полиморфизмов расположены гены, ответственные за метаболизм жирных кислот и липидов (SLC27A6), а также связанные с молочной продуктивностью и иммунным ответом (GPX8, CDC20B и GZMA). Выявленные ОНП и локусы могут выступать в качестве кандидатных маркеров для геномной селекции голштинского скота. Совпадение GWAS-результатов на полной выборке и экстремальных квартилях Q1/Q4 подтверждает надежность выявленных ассоциаций и эффективность использования метода экстремальных выборок.

The aim of the study is to identify genetic markers associated with dairy productivity parameters (milk yield, milk fat and protein content) in Holstein cattle using the genome-wide association analysis (GWAS) method on a sample of animals from breeding farms in the Sverdlovsk Region for 2018–2024. Objectives: to genotype a sample of cows using DNA bioarrays and carry out data quality control; to perform GWAS analysis of dairy productivity on the full population and on extreme phenotypic groups; to identify statistically significant single nucleotide polymorphisms (SNPs) associated with milk yield, fat and protein content; to analyze genes located near significant SNPs to understand their role in lipid metabolism, immune response and mammary tissue development; to evaluate the coincidence of genetic associations between the full sample and extreme groups to confirm the reliability of markers. GWAS analysis of milk productivity was conducted on 539 Holstein cows from 3 breeding farms in the Sverdlovsk Region (2018–2024). Milk yield, fat, and protein were analyzed on the full sample and at the extreme quartiles Q1/Q4 (milk yield: (10,028 ± 680) and (6,775 ± 653) kg; fat: (4.07 ± 0.068) and (3.79 ± 0.092) %; protein: (3.38 ± 0.066) and (3.12 ± 0.045) %). A total of 20 significant SNPs associated with cattle productivity parameters were identified. Genes responsible for fatty acid and lipid metabolism (SLC27A6), as well as those associated with milk productivity and the immune response (GPX8, CDC20B, and GZMA) are located near some polymorphisms. The identified SNPs and loci can serve as candidate markers for genomic selection of Holstein cattle. The agreement between the GWAS results for the full sample and the extreme quartiles Q1/Q4 confirms the reliability of the identified associations and the effectiveness of the extreme sampling method.

 GWAS молочная продуктивность ОНП геномная селекция крупный рогатый скот голштинская порода GWAS milk productivity SNPs genomic selection cattle Holstein breed исследование проведено в рамках госзадания Министерства сельского хозяйства Российской Федерации (регистрационный номер 124061300024-1). the study was conducted as part of a state assignment from the Ministry of Agriculture of the Russian Federation (registration number 124061300024-1).

Brito L.F., Bédère N., Douhard F., et al. Genetic selection of high-yielding dairy cattle toward sustainable farming systems in a rapidly changing world // Animal. 2021. Vol. 15. P. 100292. DOI: 10.1017/S175173112100056X. Brito LF, Bédère N, Douhard F, et al. Genetic selection of high-yielding dairy cattle toward sustainable farming systems in a rapidly changing world. Animal. 2021;15:100292. DOI: 10.1017/S175173112 100056X. García-Ruiz A., Cole J.B., VanRaden P.M., et al. Changes in genetic selection differentials and generation intervals in US Holstein dairy cattle as a result of genomic selection // Proceedings of the National Academy of Sciences. 2016. Vol. 113, N 28. DOI: 10.1073/pnas.1603031113. García-Ruiz A, Cole JB, VanRaden PM, et al. Changes in genetic selection differentials and generation intervals in US Holstein dairy cattle as a result of genomic selection. Proc Natl Acad Sci USA. 2016;113(28). DOI: 10.1073/pnas.1603031113. Petrov A.F., Bogdanova O.V., Narozhnykh K.N., et al. Clustering of countries based on dairy productivity characteristics of Holstein cattle for breeding material selection // Veterinary World. 2024. P. 1108–1118. Petrov AF, Bogdanova OV, Narozhnykh KN, et al. Clustering of countries based on dairy productivity characteristics of Holstein cattle for breeding material selection. Vet World. 2024:1108-1118. Dotsev A.V., Sermyagin A.A. PSXIV-4 Breed purity of Holstein bulls born in Russia and imported from different countries // Journal of Animal Science. 2018. Vol. 96, is. 3. P. 142–143. Dotsev AV, Sermyagin AA. PSXIV-4 Breed purity of Holstein bulls born in Russia and imported from different countries. J Anim Sci. 2018;96(3):142-143. Фирсова Е.В., Карташова А.П. Голштинская порода скота в Российской Федерации: современное состояние и перспективы развития // Генетика и разведение животных. 2019. № 1. С. 62–69. Firsova EV, Kartashova AP. Holstein breed of the cattle in the Russian Federation, the current state and the prospects of development. Genetics and breeding of animals. 2019;1:62-69. (In Russ.). Wang J., Daetwyler H.D., Zhang Q., et al. Genome-wide association study and fine-mapping using imputed sequences to prioritize candidate genes for 30 complex traits in 50,309 Holstein bulls // Journal of Dairy Science. 2025. Wang J, Daetwyler HD, Zhang Q, et al. Genome-wide association study and fine-mapping using imputed sequences to prioritize candidate genes for 30 complex traits in 50,309 Holstein bulls. J Dairy Sci. 2025. Ahmed R.H., Steyn Y., Banga C.B., et al. Genomic-based genetic parameters and genome-wide association studies for productive and reproductive traits in Beef-on-Dairy crossbreds // Frontiers in Genetics. 2025. Vol. 16. Ahmed RH, Steyn Y, Banga CB, et al. Genomic-based genetic parameters and genome-wide association studies for productive and reproductive traits in Beef-on-Dairy crossbreds. Front Genet. 2025;16. Lopdell T.J. Using QTL to Identify Genes and Pathways Underlying the Regulation and Production of Milk Components in Cattle // Animals. 2023. Vol. 13, N 5. P. 911. DOI: 10.3390/ani13050911. Lopdell TJ. Using QTL to Identify Genes and Pathways Underlying the Regulation and Production of Milk Components in Cattle. Animals. 2023;13(5):911. DOI: 10.3390/ani13050911. Tarsani E., Freidin S., Mason W., et al. Genome-wide association studies of dairy cattle resistance to digital dermatitis recorded at four distinct lactation stages // Scientific Reports. 2025. Vol. 15, N 1. P. 8922. DOI: 10.1038/s41598-025-13443-1. Tarsani E, Freidin S, Mason W, et al. Genome-wide association studies of dairy cattle resistance to digital dermatitis recorded at four distinct lactation stages. Sci Rep. 2025;15(1):8922. DOI: 10.1038/ s41598-025-13443-1. Wiggans G.R., Cole J.B., Null D.J., et al. Genomic Selection in Dairy Cattle: The USDA Experience // Annual Review of Animal Biosciences. 2017. Vol. 5, N 1. P. 309–327. DOI: 10.1146/annurev-animal-022516-022747. Wiggans GR, Cole JB, Null DJ, et al. Genomic Selection in Dairy Cattle: The USDA Experience. Annu Rev Anim Biosci. 2017;5(1):309-327. DOI: 10.1146/annurev-animal-022516-022747. Li D., Lewinger J.P., Gauderman W.J., et al. Using extreme phenotype sampling to identify the rare causal variants of quantitative traits in association studies // Genetic Epidemiology. 2011. Vol. 35, N 8. P. 790–799. DOI: 10.1002/gepi.20652. Li D, Lewinger JP, Gauderman WJ, et al. Using extreme phenotype sampling to identify the rare causal variants of quantitative traits in association studies. Genet Epidemiol. 2011;35(8):790-799. DOI: 10.1002/gepi.20652. Gage J.L., de Leon N., Clayton M.K. Comparing Genome-Wide Association Study Results from Different Measurements of an Underlying Phenotype // G3 Genes|Genomes|Genetics. 2018. Vol. 8, N 11. P. 3715–3722. DOI: 10.1534/g3.118.200596. Gage JL, de Leon N, Clayton MK. Comparing Genome-Wide Association Study Results from Different Measurements of an Underlying Phenotype. G3 (Bethesda). 2018;8(11):3715-3722. DOI: 10.1534/ g3.118.200596. Yang J., Loos R.J.F. Extreme-phenotype genome-wide association study (XP-GWAS): a method for identifying trait-associated variants by sequencing pools of individuals selected from a diversity panel // The Plant Journal. 2015. Vol. 84, N 3. P. 587–596. DOI: 10.1111/tpj.13031. Yang J, Loos RJF. Extreme-phenotype genome-wide association study (XP-GWAS): a method for identifying trait-associated variants by sequencing pools of individuals selected from a diversity panel. Plant J. 2015;84(3):587-596. DOI: 10.1111/tpj.13031. Lu D., Wang Z., Sargolzaei M., et al. Genome-wide association analyses for growth and feed efficiency traits in beef cattle // Journal of Animal Science. 2013. Vol. 91, N 8. P. 3612–3633. DOI: 10.2527/jas.2012-5972. Lu D, Wang Z, Sargolzaei M, et al. Genome-wide association analyses for growth and feed efficiency traits in beef cattle. J Anim Sci. 2013;91(8):3612-3633. DOI: 10.2527/jas.2012-5972. Mancin E., Tagliapietra F., Shtylla B., et al. Genome Wide Association Study of Beef Traits in Local Alpine Breed Reveals the Diversity of the Pathways Involved and the Role of Time Stratification // Frontiers in Genetics. 2022. Vol. 12. DOI: 10.3389/fgene.2021.701301. Mancin E, Tagliapietra F, Shtylla B, et al. Genome Wide Association Study of Beef Traits in Local Alpine Breed Reveals the Diversity of the Pathways Involved and the Role of Time Stratification. Front Genet. 2022;12. DOI: 10.3389/fgene.2021.701301. Zhuang Z., Luo Y., Cai Y., et al. Weighted Single-Step Genome-Wide Association Study for Growth Traits in Chinese Simmental Beef Cattle // Genes. 2020. Vol. 11, N 2. P. 189. DOI: 10.3390/genes11020189. Zhuang Z, Luo Y, Cai Y, et al. Weighted Single-Step Genome-Wide Association Study for Growth Traits in Chinese Simmental Beef Cattle. Genes (Basel). 2020;11(2):189. DOI: 10.3390/genes11020189. Suzuki T., Terasaki M., Takemoto Y., et al. Structural Compensation for the Deficit of rRNA with Proteins in the Mammalian Mitochondrial Ribosome // Journal of Biological Chemistry. 2001. Vol. 276, N 24. P. 21724–21736. DOI: 10.1074/jbc.M101787200. Suzuki T, Terasaki M, Takemoto Y, et al. Structural Compensation for the Deficit of rRNA with Proteins in the Mammalian Mitochondrial Ribosome. J Biol Chem. 2001;276(24):21724-21736. DOI: 10.1074/jbc.M101787200. МерckKGaA. COLEC10. Доступно по: https://sigmaaldrich.com/NL/en/genes/colec10. Ссылка активна на 25.12.2025. Merck KGaA. COLEC10. Available at: https://sigmaaldrich.com/NL/en/genes/colec10. Accessed 25.12.2025. Sharma A., Prowse-Wilkins C., Reverter A., et al. Stories and Challenges of Genome Wide Association Studies in Livestock – A Review // Asian-Australasian Journal of Animal Sciences. 2015. Vol. 28, N 10. P. 1371–1379. DOI: 10.5713/ajas.15.0388. Sharma A, Prowse-Wilkins C, Reverter A, et al. Stories and Challenges of Genome Wide Association Studies in Livestock – A Review. Asian-Australas J Anim Sci. 2015;28(10):1371-1379. DOI: 10.5713/ajas.15.0388. Bang N.N., Haile-Mariam M., Pryce J.E., et al. Genomic Prediction and Genome‐Wide Association Studies for Productivity, Conformation and Heat Tolerance Traits in Tropical Smallholder Dairy Cows // Journal of Animal Breeding and Genetics. 2025. Vol. 142, N 3. P. 322–341. DOI: 10.1111/jbg.12532. Bang NN, Haile-Mariam M, Pryce JE, et al. Genomic Prediction and Genome‐Wide Association Studies for Productivity, Conformation and Heat Tolerance Traits in Tropical Smallholder Dairy Cows. J Anim Breed Genet. 2025;142(3):322-341. DOI: 10.1111/jbg.12532. Meredith B.K., Berry D.P., Pryce J.E., et al. Genome-wide associations for milk production and somatic cell score in Holstein-Friesian cattle in Ireland // BMC Genetics. 2012. Vol. 13, N 1. P. 21. DOI: 10.1186/1471-2156-13-21. Meredith BK, Berry DP, Pryce JE, et al. Genome-wide associations for milk production and somatic cell score in Holstein-Friesian cattle in Ireland. BMC Genet. 2012;13:21. DOI: 10.1186/1471-2156-13-21. Azumah R., Lutsay V., Merriman J.A., et al. Candidate genes for polycystic ovary syndrome are regulated by TGFβ in the bovine foetal ovary // Human Reproduction. 2022. Vol. 37, N 6. P. 1244–1254. DOI: 10.1093/humrep/deac062. Azumah R, Lutsay V, Merriman JA, et al. Candidate genes for polycystic ovary syndrome are regulated by TGFβ in the bovine foetal ovary. Hum Reprod. 2022;37(6):1244-1254. DOI: 10.1093/humrep/deac062. Bastian N.A., Humpherson P.G., Lane E.B., et al. Regulation of fibrillins and modulators of TGFβ in fetal bovine and human ovaries // Reproduction. 2016. Vol. 152, N 2. P. 127–137. DOI: 10.1530/REP-15-0396. Bastian NA, Humpherson PG, Lane EB, et al. Regulation of fibrillins and modulators of TGFβ in fetal bovine and human ovaries. Reproduction. 2016;152(2):127-137. DOI: 10.1530/REP-15-0396. Gao J., Cheng Y., Niu C., et al. Evidence of early genomic selection in Holstein Friesian across African and European ecosystems // BMC Genomics. 2025. Vol. 26, N 1. P. 615. DOI: 10.1186/s12864-025-19095-5. Gao J, Cheng Y, Niu C, et al. Evidence of early genomic selection in Holstein Friesian across African and European ecosystems. BMC Genomics. 2025;26(1):615. DOI: 10.1186/s12864-025-19095-5. Wang X., Huang Y., Zhai Y., et al. Identification of the hub genes related to adipose tissue metabolism of bovine // Frontiers in Veterinary Science. 2022. Vol. 9. DOI: 10.3389/fvets.2022.832197. Wang X, Huang Y, Zhai Y, et al. Identification of the hub genes related to adipose tissue metabolism of bovine. Front Vet Sci. 2022;9. DOI: 10.3389/fvets.2022.832197. Ren E., Wang Y., Zhou B., et al. Transcriptomic and metabolomic responses induced in the livers of growing pigs by a short-term intravenous infusion of sodium butyrate // Animal. 2018. Vol. 12, N 11. P. 2318–2326. DOI: 10.1017/S1751731118000283. Ren E, Wang Y, Zhou B, et al. Transcriptomic and metabolomic responses induced in the livers of growing pigs by a short-term intravenous infusion of sodium butyrate. Animal. 2018;12(11):2318-2326. DOI: 10.1017/S1751731118000283. Peterson T.R., Laplante M., Thoreen C.C., et al. DEPTOR Is an mTOR Inhibitor Frequently Overexpressed in Multiple Myeloma Cells and Required for Their Survival // Cell. 2009. Vol. 137, N 5. P. 873–886. DOI: 10.1016/j.cell.2009.02.024. Peterson TR, Laplante M, Thoreen CC, et al. DEPTOR Is an mTOR Inhibitor Frequently Overexpressed in Multiple Myeloma Cells and Required for Their Survival. Cell. 2009;137(5):873-886. DOI: 10.1016/j.cell.2009.02.024. Agarwal N., Santhoshkumar T., Shukla A.K., et al. MTBP plays a crucial role in mitotic progression and chromosome segregation // Cell Death & Differentiation. 2011. Vol. 18, N 7. P. 1208–1219. DOI: 10.1038/cdd.2011.22. Agarwal N, Santhoshkumar T, Shukla AK, et al. MTBP plays a crucial role in mitotic progression and chromosome segregation. Cell Death Differ. 2011;18(7):1208-1219. DOI: 10.1038/cdd.2011.22. Metzger J., Chen S., Willis S., et al. Whole genome sequencing identifies missense mutation in MTBP in Shar-Pei affected with Autoinflammatory Disease (SPAID) // BMC Genomics. 2017. Vol. 18, N 1. P. 348. DOI: 10.1186/s12864-017-3746-2. Metzger J, Chen S, Willis S, et al. Whole genome sequencing identifies missense mutation in MTBP in Shar-Pei affected with Autoinflammatory Disease (SPAID). BMC Genomics. 2017;18(1):348. DOI: 10.1186/s12864-017-3746-2. Zhang Y., Zhao S., Wei Q., et al. Acetyl‐coenzyme A acyltransferase 2 promote the differentiation of sheep precursor adipocytes into adipocytes // Journal of Cellular Biochemistry. 2019. Vol. 120, N 5. P. 8021–8031. DOI: 10.1002/jcb.27908. Zhang Y, Zhao S, Wei Q, et al. Acetyl‐coenzyme A acyltransferase 2 promotes the differentiation of sheep precursor adipocytes into adipocytes. J Cell Biochem. 2019;120(5):8021-8031. DOI: 10.1002/ jcb.27908. Durán Aguilar M., Gutiérrez-Gil B., Arranz J.J., et al. Genome‐wide association study for milk somatic cell score in holsteincattle using copy number variation as markers // Journal of Animal Breeding and Genetics. 2017. Vol. 134, N 1. P. 49–59. DOI: 10.1111/jbg.12217. Durán Aguilar M, Gutiérrez-Gil B, Arranz JJ, et al. Genome‐wide association study for milk somatic cell score in holstein cattle using copy number variation as markers. J Anim Breed Genet. 2017;134(1):49-59. DOI: 10.1111/jbg.12217. Wang G., Pan T., Jiang H., et al. Characterization of PRDM9 Multifunctionality in Yak Testes Through Protein Interaction Mapping // International Journal of Molecular Sciences. 2025. Vol. 26, N 4. P. 1420. DOI: 10.3390/ijms26041420. Wang G, Pan T, Jiang H, et al. Characterization of PRDM9 Multifunctionality in Yak Testes Through Protein Interaction Mapping. Int J Mol Sci. 2025;26(4):1420. DOI: 10.3390/ijms26041420. Gai Z., Hu X., Chen Y., et al. L-arginine alleviates heat stress-induced mammary gland injury through modulating CASTOR1-mTORC1 axis mediated mitochondrial homeostasis // Science of The Total Environment. 2024. Vol. 926. P. 172017. DOI: 10.1016/j.scitotenv.2023.172017. Gai Z, Hu X, Chen Y, et al. L-arginine alleviates heat stress-induced mammary gland injury through modulating CASTOR1-mTORC1 axis mediated mitochondrial homeostasis. Sci Total Environ. 2024;926:172017. DOI: 10.1016/j.scitotenv.2023.172017. Otto P.I., Botha A., de Kock H.L., et al. Genome-wide association studies for heat stress response in Bos taurus × Bos indicus crossbred cattle // Journal of Dairy Science. 2019. Vol. 102, N 9. P. 8148–8158. DOI: 10.3168/jds.2018-15676. Otto PI, Botha A, de Kock HL, et al. Genome-wide association studies for heat stress response in Bos taurus × Bos indicus crossbred cattle. J Dairy Sci. 2019;102(9):8148-8158. DOI: 10.3168/jds.2018-15676. Casarotto L.T., Garcia-Ruiz A.L., Lima F.S., et al. Late-gestation heat stress alters placental structure and function in multiparous dairy cows // Journal of Dairy Science. 2025. Vol. 108, N 1. P. 1125–1137. DOI: 10.3168/jds.2024-23707. Casarotto LT, Garcia-Ruiz AL, Lima FS, et al. Late-gestation heat stress alters placental structure and function in multiparous dairy cows. J Dairy Sci. 2025;108(1):1125-1137. DOI: 10.3168/jds.2024-23707. Nayak S.S., Kumar S., Verma K., et al. Deciphering climate resilience in Indian cattle breeds by selection signature analyses // Tropical Animal Health and Production. 2024. Vol. 56, N 2. P. 46. DOI: 10.1007/s11250-023-03624-6. Nayak SS, Kumar S, Verma K, et al. Deciphering climate resilience in Indian cattle breeds by selection signature analyses. Trop Anim Health Prod. 2024;56(2):46. DOI: 10.1007/s11250-023-03624-6. Guo Y., Wang L., Wang Y., et al. Comparative Transcriptome Analysis of Bovine, Porcine, and Sheep Muscle Using Interpretable Machine Learning Models // Animals. 2024. Vol. 14, N 20. P. 2947. DOI: 10.3390/ani14202947. Guo Y, Wang L, Wang Y, et al. Comparative Transcriptome Analysis of Bovine, Porcine, and Sheep Muscle Using Interpretable Machine Learning Models. Animals (Basel). 2024;14(20):2947. DOI: 10.3390/ani14202947. Sun R., Zhang Q., Wang X., et al. Molecular Characterization, Expression Profiles of SMAD4, SMAD5 and SMAD7 Genes and Lack of Association with Litter Size in Tibetan Sheep // Animals. 2022. Vol. 12, N 17. P. 2232. DOI: 10.3390/ani12172232. Sun R, Zhang Q, Wang X, et al. Molecular Characterization, Expression Profiles of SMAD4, SMAD5 and SMAD7 Genes and Lack of Association with Litter Size in Tibetan Sheep. Animals (Basel). 2022;12(17):2232. DOI: 10.3390/ani12172232. Wei D., Yang Y., Wang S., et al. Roles of MEF2A and HOXA5 in the transcriptional regulation of the bovine Fox O1 gene // Animal Biotechnology. 2023. Vol. 34, N 9. P. 4367–4379. DOI: 10.1080/ 10495398.2022.2106683. Wei D, Yang Y, Wang S, et al. Roles of MEF2A and HOXA5 in the transcriptional regulation of the bovine Fox O1 gene. Anim Biotechnol. 2023;34(9):4367-4379. DOI: 10.1080/10495398.2022.2106683. Li P., Wang H., Li Q., et al. Multi-omics analysis of lipids and aroma compounds in beef under grain-fed and grass-fed production methods // Current Research in Food Science. 2025. Vol. 11. P. 101216. DOI: 10.1016/j.crfs.2024.101216. Li P, Wang H, Li Q, et al. Multi-omics analysis of lipids and aroma compounds in beef under grain-fed and grass-fed production methods. Curr Res Food Sci. 2025;11:101216. DOI: 10.1016/j.crfs.2024. 101216. Yu H., Chen L., Zhang K., et al. Integrated multi-omics analysis reveals variation in intramuscular fat among muscle locations of Qinchuan cattle // BMC Genomics. 2023. Т. 24, № 1. С. 367. DOI: 10.1186/s12864-023-09487-9. Yu H, Chen L, Zhang K, et al. Integrated multi-omics analysis reveals variation in intramuscular fat among muscle locations of Qinchuan cattle. BMC Genomics. 2023;24(1):367. DOI: 10.1186/s12864-023-09487-9. Hosseinzadeh S., Rafat S.A., Fang L. Integrated TWAS, GWAS, and RNAseq results identify candidate genes associated with reproductive traits in cows // Scientific Reports. 2025. Т. 15, № 1. С. 1932. DOI: 10.1038/s41598-025-16487-4. Hosseinzadeh S, Rafat SA, Fang L. Integrated TWAS, GWAS, and RNAseq results identify candidate genes associated with reproductive traits in cows. Sci Rep. 2025;15(1):1932. DOI: 10.1038/s41598-025-16487-4. Jiang W., Mooney M.H., Shirali M. Unveiling the Genetic Landscape of Feed Efficiency in Holstein Dairy Cows: Insights into Heritability, Genetic Markers, and Pathways via Meta-Analysis // Journal of Animal Science. 2024. Т. 102. DOI: 10.1093/jas/skad026. Jiang W, Mooney MH, Shirali M. Unveiling the Genetic Landscape of Feed Efficiency in Holstein Dairy Cows: Insights into Heritability, Genetic Markers, and Pathways via Meta-Analysis. J Anim Sci. 2024;102. DOI: 10.1093/jas/skad026. Zheng W., Jiang J., Li X., et al. A pilot multi-omics study reveals genetic mechanisms regulating milk component traits in dairy cattle // Communications Biology. 2025. Т. 8, № 1. С. 1150. DOI: 10.1038/ s42003-025-02110-1. Zheng W, Jiang J, Li X, et al. A pilot multi-omics study reveals genetic mechanisms regulating milk component traits in dairy cattle. Commun Biol. 2025;8(1):1150. DOI: 10.1038/s42003-025-02110-1. Gebreyesus G., Zhang H., Jiang L., et al. Vitamin B12 and transcobalamin in bovine milk: Genetic variation and genome-wide association with loci along the genome // JDS Communications. 2021. Т. 2, № 3. С. 127–131. DOI: 10.3168/jdsc.2021-0108. Gebreyesus G, Zhang H, Jiang L, et al. Vitamin B12 and transcobalamin in bovine milk: genetic variation and genome-wide association with loci along the genome. JDS Commun. 2021;2(3):127-131. DOI: 10.3168/jdsc.2021-0108.