{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,29]],"date-time":"2026-09-29T23:12:29Z","timestamp":1790723549072,"version":"4.1.0"},"reference-count":90,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2026,9,4]],"date-time":"2026-09-04T00:00:00Z","timestamp":1788480000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Research and Integration of Production Technology for Special Tibetan Sheep in High Saline and Alkaline Habita","award":["2022-NK-169-3"],"award-info":[{"award-number":["2022-NK-169-3"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Metabolites"],"abstract":"<jats:p>Background: Intramuscular fat (IMF) is a key determinant of meat sensory quality, but the effects of rumen-protected betaine (RPB) on IMF deposition in Tibetan sheep remain unclear. Methods: Sixty male, 3-month-old Tibetan lambs (Initial weight 17.72 \u00b1 1.36 kg) were randomly assigned to a control (basal diet, n = 30) or RPB supplementation (basal diet supplemented with 0.08% RPB, n = 30) for 90 d. Results: Transcriptomic and lipidomic analyses revealed that RPB modulated glycerophospholipid metabolism-related genes (DGKI and AGPAT1) and altered lipid profiles, including PC (16:0\/20:5) and PC (16:0\/16:0). RPB increased T-AOC and SOD activities and decreased MDA content (p &lt; 0.05). Intramuscular adipocyte area and diameter increased, while density decreased (p &lt; 0.05). LC-MS\/MS analysis showed that unsaturated fatty acids C14:1, C17:1, and C22:4 were significantly increased (p &lt; 0.05). Muscle fiber density in the longissimus dorsi increased, whereas shear force, cooking loss, hardness, and chewiness were reduced (p &lt; 0.05). MYH1, MYH2, and MYH7 expression were upregulated by 2.05-, 1.89-, and 2.73-fold, respectively. Conclusions: In summary, dietary RPB supplementation was associated with increased IMF deposition, which may contribute to improved meat quality, potentially through modulation of glycerophospholipid metabolism.<\/jats:p>","DOI":"10.3390\/metabo16090650","type":"journal-article","created":{"date-parts":[[2026,9,4]],"date-time":"2026-09-04T12:20:46Z","timestamp":1788524446000},"page":"650","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Rumen-Protected Betaine Improves Meat Quality in Tibetan Sheep by Influencing Intramuscular Fat Deposition, Antioxidant Capacity, and Fatty Acid Composition"],"prefix":"10.3390","volume":"16","author":[{"ORCID":"https:\/\/orcid.org\/0009-0006-8957-6592","authenticated-orcid":false,"given":"Chengdi","family":"Shi","sequence":"first","affiliation":[{"name":"College of Agriculture and Animal Husbandry, Qinghai University, Xining City 810016, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"data-curation","vocabulary":"credit"},{"role":"conceptualization","vocabulary":"credit"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"methodology","vocabulary":"credit"},{"role":"software","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"}]},{"given":"Wei","family":"Gao","sequence":"additional","affiliation":[{"name":"College of Agriculture and Animal Husbandry, Qinghai University, Xining City 810016, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"conceptualization","vocabulary":"credit"},{"role":"data-curation","vocabulary":"credit"}]},{"given":"Zhenglu","family":"Yang","sequence":"additional","affiliation":[{"name":"College of Agriculture and Animal Husbandry, Qinghai University, Xining City 810016, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"methodology","vocabulary":"credit"},{"role":"validation","vocabulary":"credit"}]},{"given":"Lijuan","family":"Han","sequence":"additional","affiliation":[{"name":"College of Agriculture and Animal Husbandry, Qinghai University, Xining City 810016, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"writing-review-editing","vocabulary":"credit"},{"role":"resources","vocabulary":"credit"}]},{"given":"Shengzhen","family":"Hou","sequence":"additional","affiliation":[{"name":"College of Agriculture and Animal Husbandry, Qinghai University, Xining City 810016, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"validation","vocabulary":"credit"},{"role":"resources","vocabulary":"credit"}]},{"given":"Chao","family":"Yang","sequence":"additional","affiliation":[{"name":"College of Agriculture and Animal Husbandry, Qinghai University, Xining City 810016, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"resources","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"},{"role":"formal-analysis","vocabulary":"credit"}]},{"given":"Zenghai","family":"Luo","sequence":"additional","affiliation":[{"name":"Qinghai Agri-Animal Husbandry Vocational College, Xining City 812100, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"investigation","vocabulary":"credit"},{"role":"validation","vocabulary":"credit"},{"role":"resources","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6392-3513","authenticated-orcid":false,"given":"Linsheng","family":"Gui","sequence":"additional","affiliation":[{"name":"College of Agriculture and Animal Husbandry, Qinghai University, Xining City 810016, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"resources","vocabulary":"credit"},{"role":"validation","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"},{"role":"funding-acquisition","vocabulary":"credit"}]}],"member":"1968","published-online":{"date-parts":[[2026,9,4]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"539","DOI":"10.1093\/ajcn\/80.3.539","article-title":"Betaine in human nutrition","volume":"80","author":"Craig","year":"2004","journal-title":"Am. J. Clin. Nutr."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"666","DOI":"10.1271\/bbb.120839","article-title":"The Effect of a Continuous Supply of Betaine on the Degradation of Betaine in the Rumen of Dairy Cows","volume":"77","author":"Nakai","year":"2013","journal-title":"Biosci. Biotechnol. Biochem."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"764","DOI":"10.2527\/jas1979.493764x","article-title":"Metabolism of Betaine in the Ruminant","volume":"49","author":"Mitchell","year":"1979","journal-title":"J. Anim. Sci."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"109920","DOI":"10.1016\/j.meatsci.2025.109920","article-title":"Effect of rumen-protected betaine and choline on carcass characteristics, amino acid and fatty acid composition of Ao-hu sheep","volume":"229","author":"Li","year":"2025","journal-title":"Meat Sci."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"1167","DOI":"10.1007\/s00394-021-02738-2","article-title":"Betaine in ameliorating alcohol-induced hepatic steatosis","volume":"61","author":"Rehman","year":"2022","journal-title":"Eur. J. Nutr."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"2792","DOI":"10.1007\/s00018-006-6249-6","article-title":"Betaine homocysteine S-methyltransferase: Just a regulator of homocysteine metabolism?","volume":"63","author":"Pajares","year":"2006","journal-title":"Cell. Mol. Life Sci. CMLS"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"7577","DOI":"10.4238\/2014.September.12.25","article-title":"Effects of dietary betaine supplementation subjected to heat stress on milk performances and physiology indices in dairy cow","volume":"13","author":"Zhang","year":"2014","journal-title":"Genet. Mol. Res."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"3363","DOI":"10.1007\/s00484-025-03026-y","article-title":"Dose-dependent effects of rumen-protected betaine on growth performance, thermoregulation, antioxidant status, and metabolic parameters in heat-stressed lambs","volume":"69","author":"Bokharaeian","year":"2025","journal-title":"Int. J. Biometeorol."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"116576","DOI":"10.1016\/j.anifeedsci.2025.116576","article-title":"Effects of rumen-protected betaine on performance, rumen function, nutrient digestibility, and nitrogen metabolism in heat-stressed lambs","volume":"331","author":"Bokharaeian","year":"2026","journal-title":"Anim. Feed. Sci. Technol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"435","DOI":"10.1017\/S1751731119002258","article-title":"Effects of rumen-protected betaine supplementation on meat quality and the composition of fatty and amino acids in growing lambs","volume":"14","author":"Dong","year":"2020","journal-title":"Animal"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"150579","DOI":"10.1016\/j.foodchem.2026.150579","article-title":"Betaine improves lamb meat quality with oxidative muscle fiber remodeling and enhanced amino acid-derived taste potential","volume":"526","author":"Zhang","year":"2026","journal-title":"Food Chem."},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Wang, Y.-Y., Zhang, Q., Han, S., Li, Y., Wang, B., Yuan, G., Zhang, P., Yang, Z.-W., Zhang, H., and Sun, Y. (2021). Ehrlichia chaffeensis and Four Anaplasma Species with Veterinary and Public Health Significance Identified in Tibetan Sheep (Ovis aries) and Yaks (Bos grunniens) in Qinghai, China. Front. Vet. Sci., 8.","DOI":"10.3389\/fvets.2021.727166"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"136666","DOI":"10.1016\/j.foodchem.2023.136666","article-title":"Identification of muttony-related compounds in cooked mutton tallows and their flavor intensities subjected to phenolic extract from thyme (Thymus vulgaris L.)","volume":"427","author":"Wu","year":"2023","journal-title":"Food Chem."},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Shi, S., Li, X., Ma, S., Sha, Y., Qu, Y., and Zhao, S. (2026). Coated Betaine Improves Lamb Meat Quality and Flavor by Modulating Rumen Microbial Flora. Animals, 16.","DOI":"10.3390\/ani16060970"},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Wang, S., Tang, W., Jiang, T., Wang, R., Zhang, R., Ou, J., Wang, Q., Cheng, X., Ren, C., and Chen, J. (2024). Effect of Dietary Concentrate-to-Forage Ratios During the Cold Season on Slaughter Performance, Meat Quality, Rumen Fermentation and Gut Microbiota of Tibetan Sheep. Animals, 14.","DOI":"10.3390\/ani14223305"},{"key":"ref_16","unstructured":"(2006). Nutrient Requirements of Meat Sheep (Standard No. NY\/T 816-2021)."},{"key":"#cr-split#-ref_17.1","unstructured":"European Commission (2018). Commission Implementing Regulation"},{"key":"#cr-split#-ref_17.2","unstructured":"(EU) 2018\/1039. Off. J. Eur. Union, L 186, 3."},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Percie du Sert, N., Hurst, V., Ahluwalia, A., Alam, S., Avey, M.T., Baker, M., Browne, W.J., Clark, A., Cuthill, I.C., and Dirnagl, U. (2020). The ARRIVE guidelines 2.0: Updated guidelines for reporting animal research. PLoS Biol., 18.","DOI":"10.1371\/journal.pbio.3000410"},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"Han, Y., He, X., Yun, Y., Chen, L., Huang, Y., Wu, Q., Qin, X., Wu, H., Wu, J., and Sha, R. (2024). The Characterization of Subcutaneous Adipose Tissue in Sunit Sheep at Different Growth Stages: A Comprehensive Analysis of the Morphology, Fatty Acid Profile, and Metabolite Profile. Foods, 13.","DOI":"10.3390\/foods13040544"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"497","DOI":"10.1016\/S0021-9258(18)64849-5","article-title":"A Simple Method for the Isolation and Purification of Total Lipides from Animal Tissues","volume":"226","author":"Folch","year":"1957","journal-title":"J. Biol. Chem."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1072","DOI":"10.1016\/S0022-2275(20)38081-0","article-title":"A simple procedure for rapid transmethylation of glycerolipids and cholesteryl esters","volume":"23","author":"Christie","year":"1982","journal-title":"J. Lipid Res."},{"key":"ref_22","first-page":"20205","article-title":"American Meat Science Association Research Guidelines for Cookery, Sensory Evaluation, and Instrumental Texture Measurements of Meat","volume":"10","author":"Miller","year":"2026","journal-title":"Meat Muscle Biol."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"416","DOI":"10.4314\/sajas.v51i4.1","article-title":"Carcass characteristics and meat quality of lambs fed high levels of spineless cactus in the diet","volume":"51","author":"Bezerra","year":"2021","journal-title":"S. Afr. J. Anim. Sci."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"447","DOI":"10.1016\/S0309-1740(98)00034-5","article-title":"Reference methods for the assessment of physical characteristics of meat","volume":"49","author":"Honikel","year":"1998","journal-title":"Meat Sci."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"108103","DOI":"10.1016\/j.foodcont.2021.108103","article-title":"Texture methods for evaluating meat and meat analogue structures: A review","volume":"127","author":"Schreuders","year":"2021","journal-title":"Food Control"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"1137","DOI":"10.1194\/jlr.D700041-JLR200","article-title":"Lipid extraction by methyl-tert-butyl ether for high-throughput lipidomics","volume":"49","author":"Matyash","year":"2008","journal-title":"J. Lipid Res."},{"key":"ref_27","doi-asserted-by":"crossref","unstructured":"Rodr\u00edguez, J.P., Casas, J., Balboa, M.A., and Balsinde, J. (2025). Bioactive lipid signaling and lipidomics in macrophage polarization: Impact on inflammation and immune regulation. Front. Immunol., 16.","DOI":"10.3389\/fimmu.2025.1550500"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1111\/j.2517-6161.1995.tb02031.x","article-title":"Controlling the False Discovery Rate: A Practical and Powerful Approach to Multiple Testing","volume":"57","author":"Benjamini","year":"1995","journal-title":"J. R. Stat. Soc. Ser. B (Methodol.)"},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Schroeder, A., Mueller, O., Stocker, S., Salowsky, R., Leiber, M., Gassmann, M., Lightfoot, S., Menzel, W., Granzow, M., and Ragg, T. (2006). The RIN: An RNA integrity number for assigning integrity values to RNA measurements. BMC Mol. Biol., 7.","DOI":"10.1186\/1471-2199-7-3"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"i884","DOI":"10.1093\/bioinformatics\/bty560","article-title":"fastp: An ultra-fast all-in-one FASTQ preprocessor","volume":"34","author":"Chen","year":"2018","journal-title":"Bioinformatics"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"907","DOI":"10.1038\/s41587-019-0201-4","article-title":"Graph-based genome alignment and genotyping with HISAT2 and HISAT-genotype","volume":"37","author":"Kim","year":"2019","journal-title":"Nat. Biotechnol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"923","DOI":"10.1093\/bioinformatics\/btt656","article-title":"featureCounts: An efficient general purpose program for assigning sequence reads to genomic features","volume":"30","author":"Liao","year":"2014","journal-title":"Bioinformatics"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"550","DOI":"10.1186\/s13059-014-0550-8","article-title":"Moderated estimation of fold change and dispersion for RNA-seq data with DESeq2","volume":"15","author":"Love","year":"2014","journal-title":"Genome Biol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"284","DOI":"10.1089\/omi.2011.0118","article-title":"clusterProfiler: An R package for comparing biological themes among gene clusters","volume":"16","author":"Yu","year":"2012","journal-title":"Omics"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"402","DOI":"10.1006\/meth.2001.1262","article-title":"Analysis of relative gene expression data using real-time quantitative PCR and the 2(-Delta Delta C(T)) Method","volume":"25","author":"Livak","year":"2001","journal-title":"Methods"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"591","DOI":"10.1093\/biomet\/52.3-4.591","article-title":"An Analysis of Variance Test for Normality (Complete Samples)","volume":"52","author":"Shapiro","year":"1965","journal-title":"Biometrika"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"364","DOI":"10.1080\/01621459.1974.10482955","article-title":"Robust Tests for the Equality of Variances","volume":"69","author":"Brown","year":"1974","journal-title":"J. Am. Stat. Assoc."},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"George, D., and Mallery, P. (2021). IBM SPSS Statistics 27 Step by Step: A Simple Guide and Reference, Routledge.","DOI":"10.4324\/9781003205333"},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Zeng, Y., Mou, H., He, Y., Zhang, D., Pan, X., Zhou, L., Shen, Y., and E, G. (2024). Effects of Key Rumen Bacteria and Microbial Metabolites on Fatty Acid Deposition in Goat Muscle. Animals, 14.","DOI":"10.3390\/ani14223225"},{"key":"ref_40","unstructured":"R Core Team (2024). R: A Language and Environment for Statistical Computing, R Core Team."},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Xu, F., Wang, H., Qin, C., Yue, B., Yang, Y., Wang, J., Zhong, J., and Wang, H. (2024). Combined Multi-Omics Analysis Reveals the Potential Role of ACADS in Yak Intramuscular Fat Deposition. Int. J. Mol. Sci., 25.","DOI":"10.3390\/ijms25169131"},{"key":"ref_42","doi-asserted-by":"crossref","unstructured":"Zhang, J., Zhang, Y., Wang, J., Jin, H., Qian, S., Chen, P., Wang, M., Chen, N., and Ding, L. (2023). Comparison of Antioxidant Capacity and Muscle Amino Acid and Fatty Acid Composition of Nervous and Calm Hu Sheep. Antioxidants, 12.","DOI":"10.3390\/antiox12020459"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"1112","DOI":"10.1093\/tas\/txz053","article-title":"Feeding differing direct-fed microbials and its influence on growth and haematological parameters of growing lambs","volume":"3","author":"Antwi","year":"2019","journal-title":"Transl. Anim. Sci."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"3846122","DOI":"10.1155\/2021\/3846122","article-title":"Effects of Exercise-Induced ROS on the Pathophysiological Functions of Skeletal Muscle","volume":"2021","author":"Wang","year":"2021","journal-title":"Oxidative Med. Cell. Longev."},{"key":"ref_45","first-page":"245","article-title":"Relation between fatty acid composition, vitamin e and malondialdehyde levels, and activity of antioxidant enzymes in the blood","volume":"99","author":"Beno","year":"1998","journal-title":"Bratisl. Lek. Listy"},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Duan, P., Yang, Y., Hou, L., Wu, Y., Li, J., Xu, C., and Guo, T. (2025). Residual Black Wolfberry Fruit Improves Meat Quality of Sheep by Enhancing Immune and Antioxidant Capacity. Vet. Sci., 12.","DOI":"10.3390\/vetsci12040324"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"6231","DOI":"10.1111\/ijfs.17359","article-title":"Betaine activates the Nrf2-Keap1-ARE pathway by increasing the methylation level of Keap1 DNA promoter","volume":"59","author":"Zhang","year":"2024","journal-title":"Int. J. Food Sci. Technol."},{"key":"ref_48","doi-asserted-by":"crossref","unstructured":"Ornoy, A., Becker, M., Weinstein-Fudim, L., and Ergaz, Z. (2020). S-Adenosine Methionine (SAMe) and Valproic Acid (VPA) as Epigenetic Modulators: Special Emphasis on their Interactions Affecting Nervous Tissue during Pregnancy. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21103721"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"30","DOI":"10.1186\/s13568-026-02024-6","article-title":"Rumen-protected betaine supplementation improves ileal function by affecting microbe and metabolite in Tibetan sheep","volume":"16","author":"Gao","year":"2026","journal-title":"AMB Express"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"e6","DOI":"10.1017\/anr.2024.26","article-title":"Betaine increases intramuscular fat deposition by promoting the cell cycle via a manner involving NADPH\/FTO\/m6A signaling","volume":"2","author":"Liu","year":"2025","journal-title":"Anim. Nutr."},{"key":"ref_51","doi-asserted-by":"crossref","unstructured":"de Amorim Rodrigues, G., Saraiva, A., Soares, M.H., Oliveira Mendes, T.A., Costa, T.C., Brito da Silva, C., Monteiro, P.V., and Duarte, M. (2025). Betaine supplementation affects carbohydrate metabolism in the skeletal muscle of finishing pigs. PLoS ONE, 20.","DOI":"10.1371\/journal.pone.0322040"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"739","DOI":"10.1016\/j.foodres.2016.09.010","article-title":"The structural basis of cooking loss in beef: Variations with temperature and ageing","volume":"89","author":"Purslow","year":"2016","journal-title":"Food Res. Int."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"105113","DOI":"10.1016\/j.psj.2025.105113","article-title":"Effects of environmental temperature on growth performance, muscle fiber structure, fatty acid composition, and gene expression in ducks","volume":"104","author":"Pan","year":"2025","journal-title":"Poult. Sci."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"351","DOI":"10.1016\/j.meatsci.2005.04.015","article-title":"The relationship between muscle fiber characteristics, postmortem metabolic rate, and meat quality of pig longissimus dorsi muscle","volume":"71","author":"Ryu","year":"2005","journal-title":"Meat Sci."},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Fu, Y., Chen, Y., Han, X., Tan, D., Chen, J., Lai, C., Yang, X., Shan, X., Silva, L.H.P., and Jiang, H. (2025). Effects of Muscle Fiber Composition on Meat Quality, Flavor Characteristics, and Nutritional Traits in Lamb. Foods, 14.","DOI":"10.3390\/foods14132309"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1186\/s40104-025-01279-6","article-title":"Gut microbiota and metabolites in lipid metabolism and intramuscular fat deposition: Mechanisms and implications for meat quality","volume":"16","author":"Song","year":"2025","journal-title":"J. Anim. Sci. Biotechnol."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"10450","DOI":"10.1038\/s41598-021-90012-0","article-title":"iTRAQ-mediated analysis of the relationship between proteomic changes and yak longissimus lumborum tenderness over the course of postmortem storage","volume":"11","author":"Yang","year":"2021","journal-title":"Sci. Rep."},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Li, R., Li, B., Jiang, A., Cao, Y., Hou, L., Zhang, Z., Zhang, X., Liu, H., Kim, K.-H., and Wu, W. (2020). Exploring the lncRNAs Related to Skeletal Muscle Fiber Types and Meat Quality Traits in Pigs. Genes, 11.","DOI":"10.3390\/genes11080883"},{"key":"ref_59","doi-asserted-by":"crossref","unstructured":"Pe\u0107ina, M., Konja\u010di\u0107, M., Ugarkovi\u0107, N.K., and Ivankovi\u0107, A. (2023). Effect of FASN, SCD, and GH Genes on Carcass Fatness and Fatty Acid Composition of Intramuscular Lipids in F1 Holstein \u00d7 Beef Breeds. Agriculture, 13.","DOI":"10.3390\/agriculture13030571"},{"key":"ref_60","doi-asserted-by":"crossref","unstructured":"Hu, C., Ding, L., Jiang, C., Ma, C., Liu, B., Li, D., and Degen, A.A. (2021). Effects of Management, Dietary Intake, and Genotype on Rumen Morphology, Fermentation, and Microbiota, and on Meat Quality in Yaks and Cattle. Front. Nutr., 8.","DOI":"10.3389\/fnut.2021.755255"},{"key":"ref_61","doi-asserted-by":"crossref","unstructured":"Chen, B., Yue, Y., Li, J., Liu, J., Yuan, C., Guo, T., Zhang, D., Yang, B., and Lu, Z. (2022). Transcriptome-metabolome analysis reveals how sires affect meat quality in hybrid sheep populations. Front. Nutr., 9.","DOI":"10.3389\/fnut.2022.967985"},{"key":"ref_62","doi-asserted-by":"crossref","unstructured":"Chen, H., Peng, T., Shang, H., Shang, X., Zhao, X., Qu, M., and Song, X. (2022). RNA-Seq Analysis Reveals the Potential Molecular Mechanisms of Puerarin on Intramuscular Fat Deposition in Heat-Stressed Beef Cattle. Front. Nutr., 9.","DOI":"10.3389\/fnut.2022.817557"},{"key":"ref_63","doi-asserted-by":"crossref","unstructured":"Liu, L., Cao, P., Zhang, L., Qi, M., Wang, L., Li, Z., Shao, G., Ding, L., Zhao, X., and Zhao, X. (2021). Comparisons of adipogenesis- and lipid metabolism-related gene expression levels in muscle, adipose tissue and liver from Wagyu-cross and Holstein steers. PLoS ONE, 16.","DOI":"10.1371\/journal.pone.0247559"},{"key":"ref_64","doi-asserted-by":"crossref","unstructured":"Pei, Y., Song, Y., Feng, Z., Li, H., Mu, Y., Rehman, S.U., Liu, Q., and Li, K. (2022). Myostatin Alteration in Pigs Enhances the Deposition of Long-Chain Unsaturated Fatty Acids in Subcutaneous Fat. Foods, 11.","DOI":"10.3390\/foods11091286"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"354","DOI":"10.1038\/bjc.1990.78","article-title":"Tumour necrosis factor: A cytokine with multiple biological activities","volume":"61","author":"Semenzato","year":"1990","journal-title":"Br. J. Cancer"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"3720","DOI":"10.1182\/blood-2010-07-273417","article-title":"Interleukin-1 in the pathogenesis and treatment of inflammatory diseases","volume":"117","author":"Dinarello","year":"2011","journal-title":"Blood"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"25737","DOI":"10.1074\/jbc.272.41.25737","article-title":"Purification and characterization of the human interleukin-18 receptor","volume":"272","author":"Torigoe","year":"1997","journal-title":"J. Biol. Chem."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"44953","DOI":"10.1074\/jbc.M207923200","article-title":"Role of TRAF3 and -6 in the activation of the NF-kappa B and JNK pathways by X-linked ectodermal dysplasia receptor","volume":"277","author":"Sinha","year":"2002","journal-title":"J. Biol. Chem."},{"key":"ref_69","doi-asserted-by":"crossref","unstructured":"Truong, A.D., Hong, Y., Lee, J., Lee, K., Kil, D.Y., Lillehoj, H.S., and Hong, Y.H. (2018). Interleukin-34 Regulates Th1 and Th17 Cytokine Production by Activating Multiple Signaling Pathways through CSF-1R in Chicken Cell Lines. Int. J. Mol. Sci., 19.","DOI":"10.3390\/ijms19061665"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"118","DOI":"10.1080\/21541264.2023.2294623","article-title":"Transcription factors in the development and treatment of immune disorders","volume":"16","author":"Patalano","year":"2025","journal-title":"Transcription"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"132","DOI":"10.1080\/21623945.2020.1738792","article-title":"Involvement of calcium channels in the regulation of adipogenesis","volume":"9","author":"Zhai","year":"2020","journal-title":"Adipocyte"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"168","DOI":"10.1038\/s41419-023-05683-6","article-title":"The MYC-regulated lncRNA LNROP (ENSG00000254887) enables MYC-driven cell proliferation by controlling the expression of OCT2","volume":"14","author":"Hart","year":"2023","journal-title":"Cell Death Dis."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"7322","DOI":"10.1038\/s41598-022-10907-4","article-title":"Modified lipid metabolism and cytosolic phospholipase A2 activation in mesangial cells under pro-inflammatory conditions","volume":"12","author":"Boi","year":"2022","journal-title":"Sci. Rep."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"1375","DOI":"10.1002\/med.22015","article-title":"GADD45A: With or without you","volume":"44","author":"Palomer","year":"2024","journal-title":"Med. Res. Rev."},{"key":"ref_75","unstructured":"Rousseau, M. (2024). The Innovative Role of DUSP4 in Kidney Disease: From Diabetic to End-Stage Kidney Disease, Universit\u00e9 de Sherbrooke."},{"key":"ref_76","doi-asserted-by":"crossref","unstructured":"Sakane, F., Hoshino, F., and Murakami, C. (2020). New Era of Diacylglycerol Kinase, Phosphatidic Acid and Phosphatidic Acid-Binding Protein. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21186794"},{"key":"ref_77","doi-asserted-by":"crossref","unstructured":"Ding, Y., Hou, Y., Ling, Z., Chen, Q., Xu, T., Liu, L., Yu, N., Ni, W., Ding, X., and Zhang, X. (2022). Identification of Candidate Genes and Regulatory Competitive Endogenous RNA (ceRNA) Networks Underlying Intramuscular Fat Content in Yorkshire Pigs with Extreme Fat Deposition Phenotypes. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms232012596"},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"D944","DOI":"10.2741\/Leung","article-title":"The structure and functions of human lysophosphatidic acid acyltransferases","volume":"6","author":"Leung","year":"2001","journal-title":"Front. Biosci."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"6934","DOI":"10.1038\/s41467-023-42532-8","article-title":"Genome-wide association analysis of plasma lipidome identifies 495 genetic associations","volume":"14","author":"Ottensmann","year":"2023","journal-title":"Nat. Commun."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"474","DOI":"10.1038\/s41586-025-08942-y","article-title":"PLA2G15 is a BMP hydrolase and its targeting ameliorates lysosomal disease","volume":"642","author":"Nyame","year":"2025","journal-title":"Nature"},{"key":"ref_81","doi-asserted-by":"crossref","unstructured":"Faggion, S., Bonfatti, V., and Carnier, P. (2024). Genome-Wide Association Study for Weight Loss at the End of Dry-Curing of Hams Produced from Purebred Heavy Pigs. Animals, 14.","DOI":"10.3390\/ani14131983"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"44507","DOI":"10.1074\/jbc.M207712200","article-title":"The Surface of Lipid Droplets Is a Phospholipid Monolayer with a Unique Fatty Acid Composition","volume":"277","author":"Ozeki","year":"2002","journal-title":"J. Biol. Chem."},{"key":"ref_83","doi-asserted-by":"crossref","unstructured":"Najm, A., B\u00eerc\u0103, A.C., Niculescu, A.G., Alberts, A., Grumezescu, A.M., G\u0103l\u0103\u021beanu, B., Vasile, B., Beuran, M., Gaspar, B.S., and Hudi\u021b\u0103, A. (2025). Dipalmitoylphosphatidylcholine Lipid Vesicles for Delivering HMB, NMN, and L-Leucine in Sarcopenia Therapy. Molecules, 30.","DOI":"10.3390\/molecules30071437"},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"550","DOI":"10.1038\/nature12710","article-title":"A diurnal serum lipid integrates hepatic lipogenesis and peripheral fatty acid use","volume":"502","author":"Liu","year":"2013","journal-title":"Nature"},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"12886","DOI":"10.1073\/pnas.94.24.12886","article-title":"Myc represses transcription of the growth arrest gene gas1","volume":"94","author":"Lee","year":"1997","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"102179","DOI":"10.1016\/j.jbc.2022.102179","article-title":"Polyunsaturated fatty acids promote the rapid fusion of lipid droplets in Caenorhabditis elegans","volume":"298","author":"Wang","year":"2022","journal-title":"J. Biol. Chem."},{"key":"ref_87","doi-asserted-by":"crossref","unstructured":"Hermansen, C., Siao, R., Chua, G.G., Lee, M.R.X., Thong, A., Weingarten, M., Lindley, N., and Peterson, E.C. (2025). Short-Chain Fatty Acid Utilization in Cyberlindnera jadinii for Single-Cell Protein and Odd-Chain Fatty Acid Production. Microorganisms, 13.","DOI":"10.3390\/microorganisms13071558"},{"key":"ref_88","unstructured":"Kappler, L. (2018). Method development for valid high-resolution profiling of mitochondria and Omics investigation of mitochondrial adaptions to excess energy intake and physical exercise. [Ph.D. Thesis, Universit\u00e4t T\u00fcbingen]."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"122335","DOI":"10.1016\/j.lfs.2023.122335","article-title":"PC 18:1\/18:1 mediates the anti-inflammatory effects of exercise and remodels tumor microenvironment of hepatocellular carcinoma","volume":"336","author":"Zhang","year":"2024","journal-title":"Life Sci."}],"container-title":["Metabolites"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2218-1989\/16\/9\/650\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,4]],"date-time":"2026-09-04T12:27:24Z","timestamp":1788524844000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2218-1989\/16\/9\/650"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,4]]},"references-count":90,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["metabo16090650"],"URL":"https:\/\/doi.org\/10.3390\/metabo16090650","relation":{},"ISSN":["2218-1989"],"issn-type":[{"value":"2218-1989","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,4]]}}}