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Here, we show that dairy cows with mastitis have severe inflammation, apoptosis, and oxidative stress in mammary tissue. Eupatilin (EU), a natural compound derived from Artemisia argyi, alleviated LPS-induced pathological damage in bovine mammary epithelial cells (BMECs) by enhancing the transcriptional activity of NRF2. Knockdown of NRF2 abolished the protective effects of EU against LPS-induced inflammation, apoptosis, and oxidative stress in BMECs, whereas overexpression of NRF2 did not further enhance these effects. Importantly, the protective effect of EU against LPS-induced mastitis in mice was fully reversed by pharmacological inhibition of NRF2. This work not only identifies NRF2 as a potential therapeutic target for combating mastitis but also identifies EU as a promising therapeutic agent and an effective non-antibiotic option for the prophylaxis and therapy of mastitis.<\/jats:p>","DOI":"10.1021\/acs.jafc.6c02196","type":"journal-article","created":{"date-parts":[[2026,10,2]],"date-time":"2026-10-02T07:01:28Z","timestamp":1790924488000},"page":"31568-31582","source":"Crossref","is-referenced-by-count":0,"title":["Eupatilin Attenuates\nMastitis in Dairy Cows by Activating\nthe NRF2 Pathway"],"prefix":"10.1021","volume":"74","author":[{"given":"Yiyi","family":"Zhao","sequence":"first","affiliation":[{"name":"Yunnan Agricultural University , , ,","place":["Kunming, China, 650201"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qingqing","family":"Zhou","sequence":"additional","affiliation":[{"name":"Yunnan Agricultural University , , ,","place":["Kunming, China, 650201"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xinyuan","family":"Zhang","sequence":"additional","affiliation":[{"name":"Yunnan Agricultural University , , ,","place":["Kunming, China, 650201"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gefen","family":"Yin","sequence":"additional","affiliation":[{"name":"Yunnan Agricultural University , , ,","place":["Kunming, China, 650201"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Junlong","family":"Bi","sequence":"additional","affiliation":[{"name":"Yunnan Agricultural University , , ,","place":["Kunming, China, 650201"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jie","family":"Hou","sequence":"additional","affiliation":[{"name":"Yunnan Agricultural University , , ,","place":["Kunming, China, 650201"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaobing","family":"Li","sequence":"additional","affiliation":[{"name":"Yunnan Agricultural University , , ,","place":["Kunming, China, 650201"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4377-0972","authenticated-orcid":true,"given":"Zhiyuan","family":"Fang","sequence":"additional","affiliation":[{"name":"Yunnan Agricultural University , , ,","place":["Kunming, China, 650201"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,10,2]]},"reference":[{"issue":"8","key":"2026101408090799600_cit1","doi-asserted-by":"publisher","first-page":"486","DOI":"10.1016\/j.tibtech.2009.05.004","article-title":"Mastitis detection:\ncurrent trends and future perspectives","volume":"27","author":"Viguier","year":"2009","journal-title":"Trends\nBiotechnol"},{"key":"2026101408090799600_cit2","doi-asserted-by":"publisher","first-page":"535668","DOI":"10.3389\/fpubh.2020.535668","article-title":"Microbial\nresistance movements: an overview of global public health threats\nposed by antimicrobial resistance, and how best to counter","volume":"8","author":"Dhingra","year":"2020","journal-title":"Front Public Health"},{"issue":"3","key":"2026101408090799600_cit3","doi-asserted-by":"publisher","first-page":"2102","DOI":"10.3168\/jds.2015-9487","article-title":"Udder infections with Staphylococcus\naureus, Streptococcus dysgalactiae, and Streptococcus uberis at calving\nin dairy herds with suboptimal udder health","volume":"99","author":"Lundberg","year":"2016","journal-title":"J. Dairy Sci."},{"issue":"1","key":"2026101408090799600_cit4","doi-asserted-by":"publisher","first-page":"36","DOI":"10.1186\/1751-0147-53-36","article-title":"Etiology and\nantimicrobial\nsusceptibility of udder pathogens from cases of subclinical mastitis\nin dairy cows in Sweden","volume":"53","author":"Persson","year":"2011","journal-title":"Acta Vet. Scand"},{"issue":"4","key":"2026101408090799600_cit5","doi-asserted-by":"publisher","first-page":"357","DOI":"10.1007\/s10911-011-9236-y","article-title":"Molecular\nepidemiology of mastitis pathogens of dairy cattle and comparative\nrelevance to humans","volume":"16","author":"Zadoks","year":"2011","journal-title":"J. Mammary Gland Biol.\nNeoplasia"},{"issue":"1","key":"2026101408090799600_cit6","doi-asserted-by":"publisher","first-page":"101","DOI":"10.1007\/s10911-010-9164-2","article-title":"Epigenetic regulation of milk production in dairy cows","volume":"15","author":"Singh","year":"2010","journal-title":"J. Mammary Gland Biol. 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Dairy Sci."},{"key":"2026101408090799600_cit11","doi-asserted-by":"publisher","first-page":"85","DOI":"10.1016\/j.intimp.2016.11.036","article-title":"Mangiferin inhibits\nmastitis induced by LPS via suppressing\nNF-\u03baB and NLRP3 signaling pathways","volume":"43","author":"Qu","year":"2017","journal-title":"Int.\nImmunopharmacol"},{"issue":"1","key":"2026101408090799600_cit12","doi-asserted-by":"publisher","first-page":"949","DOI":"10.1080\/13880209.2023.2220351","article-title":"Antioxidative effect of Periplaneta americana extract\non dextran sulfate sodium-induced ulcerative colitis through activation\nof the Nrf2 signal","volume":"61","author":"Wu","year":"2023","journal-title":"Pharm. 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J. Mol. Sci"},{"key":"2026101408090799600_cit23","doi-asserted-by":"publisher","first-page":"8289","DOI":"10.12659\/MSM.917406","article-title":"Eupatilin alleviates lipopolysaccharide-induced acute\nlung injury by inhibiting inflammation and oxidative stress","volume":"25","author":"Liu","year":"2019","journal-title":"Med. Sci. Monit"},{"issue":"2","key":"2026101408090799600_cit24","doi-asserted-by":"publisher","first-page":"824","DOI":"10.1016\/j.intimp.2015.08.035","article-title":"Betulin suppresses S. aureus-induced mammary gland inflammatory injury\nby regulating PPAR-\u03b3 in mice","volume":"29","author":"Guo","year":"2015","journal-title":"Int. Immunopharmacol"},{"issue":"4","key":"2026101408090799600_cit25","doi-asserted-by":"publisher","first-page":"3805","DOI":"10.3390\/ijms24043805","article-title":"Allicin Alleviated\nLPS-Induced Mastitis via the TLR4\/NF-\u03baB Signaling Pathway in\nBovine Mammary Epithelial Cells","volume":"24","author":"Che","year":"2023","journal-title":"Int. J. Mol.\nSci."},{"issue":"2","key":"2026101408090799600_cit26","doi-asserted-by":"publisher","first-page":"1682","DOI":"10.3168\/jds.2018-15439","article-title":"High expression of\ncell death-inducing DFFA-like effector\na (CIDEA) promotes milk fat content in dairy cows with clinical ketosis","volume":"102","author":"Sun","year":"2019","journal-title":"J. Dairy Sci."},{"key":"2026101408090799600_cit27","doi-asserted-by":"publisher","first-page":"366","DOI":"10.1186\/1471-2164-9-366","article-title":"Gene networks driving\nbovine milk fat synthesis during\nthe lactation cycle","volume":"9","author":"Bionaz","year":"2008","journal-title":"BMC Genomics"},{"issue":"1","key":"2026101408090799600_cit28","doi-asserted-by":"publisher","first-page":"443","DOI":"10.1021\/acs.jafc.2c06133","article-title":"Nuciferine ameliorates nonesterified fatty acid-induced\nbovine mammary epithelial cell lipid accumulation, apoptosis, and\nimpaired migration via activating LKB1\/AMPK signaling pathway","volume":"71","author":"Li","year":"2023","journal-title":"J. Agric. Food Chem."},{"issue":"5","key":"2026101408090799600_cit29","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0095985","article-title":"Effects\nof Arginine concentration on the in vitro expression of Casein and\nmTOR pathway related genes in mammary epithelial cells from dairy\ncattle","volume":"9","author":"Wang","year":"2014","journal-title":"PLoS One"},{"key":"2026101408090799600_cit30","doi-asserted-by":"publisher","first-page":"89","DOI":"10.1146\/annurev.pharmtox.46.120604.141046","article-title":"Cell survival responses to environmental\nstresses via the Keap1-Nrf2-ARE pathway","volume":"47","author":"Kensler","year":"2007","journal-title":"Annu.\nRev. Pharmacol. Toxicol"},{"issue":"Pt B","key":"2026101408090799600_cit31","doi-asserted-by":"publisher","first-page":"2375","DOI":"10.1016\/j.ijbiomac.2022.10.024","article-title":"Lentinan inhibits oxidative\nstress and alleviates LPS-induced inflammation\nand apoptosis of BMECs by activating the Nrf2 signaling pathway","volume":"222","author":"Meng","year":"2022","journal-title":"Int. J. Biol. Macromol"},{"key":"2026101408090799600_cit32","doi-asserted-by":"publisher","first-page":"6505373","DOI":"10.1155\/2019\/6505373","article-title":"UFL1 alleviates\nlipopolysaccharide-induced cell damage and inflammation via regulation\nof the TLR4\/NF-\u03baB pathway in bovine mammary epithelial cells","volume":"2019","author":"Li","year":"2019","journal-title":"Oxid. Med. Cell. Longevity"},{"issue":"12","key":"2026101408090799600_cit33","doi-asserted-by":"publisher","first-page":"1918","DOI":"10.3390\/antiox10121918","article-title":"Oxidative stress in\ndairy cows: insights into the mechanistic mode of actions and mitigating\nstrategies","volume":"10","author":"Ayemele","year":"2021","journal-title":"Antioxidants"},{"issue":"26","key":"2026101408090799600_cit34","doi-asserted-by":"publisher","first-page":"9672","DOI":"10.1080\/10408398.2023.2215865","article-title":"Antioxidants and neurodegenerative eye disease","volume":"64","author":"Semenova","year":"2024","journal-title":"Crit. Rev. Food Sci. Nutr."},{"key":"2026101408090799600_cit35","doi-asserted-by":"publisher","first-page":"1190817","DOI":"10.3389\/fphar.2023.1190817","article-title":"Landscape\nanalysis and overview of the literature on oxidative stress and pulmonary\ndiseases","volume":"14","author":"Liu","year":"2023","journal-title":"Front. 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Sin\nB"},{"issue":"2","key":"2026101408090799600_cit43","doi-asserted-by":"publisher","DOI":"10.1128\/Spectrum.00105-21","article-title":"Dietary\nsupplementation of inulin ameliorates subclinical mastitis via regulation\nof rumen microbial community and metabolites in dairy cows","volume":"9","author":"Wang","year":"2021","journal-title":"Microbiol. Spectrum"},{"issue":"8","key":"2026101408090799600_cit44","doi-asserted-by":"publisher","first-page":"3425","DOI":"10.3390\/molecules28083425","article-title":"Plant essential oils\nas a tool in the control of bovine mastitis: an update","volume":"28","author":"Caneschi","year":"2023","journal-title":"Molecules"},{"issue":"6","key":"2026101408090799600_cit45","doi-asserted-by":"publisher","first-page":"2655","DOI":"10.1002\/vms3.959","article-title":"Evaluation\nof in vitro\nantibacterial effect of essential oil and some herbal plant extract\nused against mastitis pathogens","volume":"8","author":"Arbab","year":"2022","journal-title":"Vet. Med. Sci."},{"issue":"2","key":"2026101408090799600_cit46","doi-asserted-by":"publisher","first-page":"346","DOI":"10.3390\/pharmaceutics15020346","article-title":"Copaiba\nOil-based emulsion as a natural chemotherapeutic agent for the treatment\nof bovine mastitis: in vivo studies","volume":"15","author":"Campanholi","year":"2023","journal-title":"Pharmaceutics"},{"issue":"3","key":"2026101408090799600_cit47","doi-asserted-by":"publisher","first-page":"421","DOI":"10.1016\/S0749-0720(15)30611-3","article-title":"Economics of mastitis\nand mastitis control","volume":"9","author":"DeGraves","year":"1993","journal-title":"Vet Clin North Am. Food\nAnim Pract"},{"issue":"2","key":"2026101408090799600_cit48","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0172773","article-title":"Cholesterol crystals\nenhance TLR2-\nand TLR4-mediated pro-inflammatory cytokine responses of monocytes\nto the proatherogenic oral bacterium Porphyromonas gingivalis","volume":"12","author":"K\u00f8llgaard","year":"2017","journal-title":"PLoS One"},{"issue":"12","key":"2026101408090799600_cit49","doi-asserted-by":"publisher","first-page":"1012","DOI":"10.1016\/j.tibs.2016.09.002","article-title":"Mechanism\nand regulation of NLRP3\ninflammasome activation","volume":"41","author":"He","year":"2016","journal-title":"Trends Biochem. 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