{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T09:18:20Z","timestamp":1788254300735,"version":"build-2803163510"},"reference-count":79,"publisher":"American Chemical Society (ACS)","issue":"33","license":[{"start":{"date-parts":[[2026,8,19]],"date-time":"2026-08-19T00:00:00Z","timestamp":1787097600000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-029"},{"start":{"date-parts":[[2026,8,19]],"date-time":"2026-08-19T00:00:00Z","timestamp":1787097600000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-037"},{"start":{"date-parts":[[2026,8,19]],"date-time":"2026-08-19T00:00:00Z","timestamp":1787097600000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-045"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32473114"],"award-info":[{"award-number":["32473114"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100012166","name":"National Key Research and Development Program of China","doi-asserted-by":"publisher","award":["2023YFD1801100"],"award-info":[{"award-number":["2023YFD1801100"]}],"id":[{"id":"10.13039\/501100012166","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,8,26]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Copper (Cu) supplementation for growth promotion in agriculture poses toxicological risks to animals and humans. Although Cu overload is known to trigger endoplasmic reticulum (ER) stress, the downstream cellular mechanisms driven by this stress are not fully characterized. Here, we identify the ER stress response as the primary initiator of reticulophagy (ER-phagy) in porcine liver models. Using an in vivo weaned piglet model exposed to agriculturally relevant Cu doses and an in vitro hepatocyte system, we demonstrate that Cu exposure activates UPR branches, including PERK\/eIF2\u03b1 phosphorylation, IRE1 and ATF6 proteins, and promotes GRP78\u2013FAM134B association. This activation precipitated ER-phagy, evidenced by autophagosome formation, receptor upregulation, and LC3-II conversion. Pharmacological inhibition of ER stress (4-PBA) attenuated this autophagic response, whereas induction (PR-619) potentiated it. These findings suggest that the ER stress\u2013ER-phagy axis contributes to Cu-induced hepatocellular stress, providing a mechanistic framework for reassessing Cu safety thresholds in livestock feed.<\/jats:p>","DOI":"10.1021\/acs.jafc.5c16689","type":"journal-article","created":{"date-parts":[[2026,8,19]],"date-time":"2026-08-19T13:01:41Z","timestamp":1787144501000},"page":"26655-26669","source":"Crossref","is-referenced-by-count":0,"title":["Endoplasmic Reticulum Stress Drives Copper-Induced\nReticulophagy in Porcine Hepatocytes"],"prefix":"10.1021","volume":"74","author":[{"given":"Shanglong","family":"Yang","sequence":"first","affiliation":[{"name":"South China Agricultural University , , , ,","place":["Guangzhou, Guangdong, P. R. China, 510642"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wei","family":"Chen","sequence":"additional","affiliation":[{"name":"South China Agricultural University , , , ,","place":["Guangzhou, Guangdong, P. R. China, 510642"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hao","family":"Xi","sequence":"additional","affiliation":[{"name":"South China Agricultural University , , , ,","place":["Guangzhou, Guangdong, P. R. China, 510642"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wenyue","family":"Qiu","sequence":"additional","affiliation":[{"name":"South China Agricultural University , , , ,","place":["Guangzhou, Guangdong, P. R. China, 510642"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hui","family":"Huang","sequence":"additional","affiliation":[{"name":"South China Agricultural University , , , ,","place":["Guangzhou, Guangdong, P. R. China, 510642"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shaofeng","family":"Wang","sequence":"additional","affiliation":[{"name":"South China Agricultural University , , , ,","place":["Guangzhou, Guangdong, P. R. China, 510642"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0004-8734-4830","authenticated-orcid":true,"given":"Jianzhao","family":"Liao","sequence":"additional","affiliation":[{"name":"South China Agricultural University , , , ,","place":["Guangzhou, Guangdong, P. R. China, 510642"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4663-0641","authenticated-orcid":true,"given":"Zhaoxin","family":"Tang","sequence":"additional","affiliation":[{"name":"South China Agricultural University , , , ,","place":["Guangzhou, Guangdong, P. R. China, 510642"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,8,19]]},"reference":[{"key":"2026090104482325300_cit1","doi-asserted-by":"publisher","first-page":"277","DOI":"10.1016\/j.tem.2025.06.007","article-title":"A metabolic perspective\non cuproptosis","volume":"37","author":"Li","year":"2026","journal-title":"Trends\nEndocrinol. Metab."},{"key":"2026090104482325300_cit2","doi-asserted-by":"publisher","first-page":"441","DOI":"10.1152\/physrev.00011.2024","article-title":"Mammalian\ncopper homeostasis: physiological roles and\nmolecular mechanisms","volume":"105","author":"Lutsenko","year":"2025","journal-title":"Physiol. 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Biotechnol."},{"key":"2026090104482325300_cit5","doi-asserted-by":"publisher","first-page":"120299","DOI":"10.1016\/j.jenvman.2024.120299","article-title":"The fate of\nCu and Zn along the feed-animal-excreta-effluent\ncontinuum in swine systems according to feed and effluent treatment\nstrategies","volume":"354","author":"Gourlez","year":"2024","journal-title":"J. Environ. Manage."},{"key":"2026090104482325300_cit6","doi-asserted-by":"publisher","first-page":"407","DOI":"10.17221\/501\/2023-PSE","article-title":"Copper contamination in agricultural\nsoils: a review of the effects of climate, soil properties, and prolonged\ncopper pesticide application in vineyards and orchards","volume":"70","author":"Neaman","year":"2024","journal-title":"Plant Soil Environ"},{"key":"2026090104482325300_cit7","doi-asserted-by":"publisher","DOI":"10.2903\/j.efsa.2023.7728","article-title":"Re-evaluation of the existing health-based guidance\nvalues for copper and exposure assessment from all sources","volume":"21","author":"More","year":"2023","journal-title":"EFSA J."},{"key":"2026090104482325300_cit8","doi-asserted-by":"publisher","first-page":"321","DOI":"10.1016\/j.aninu.2021.10.007","article-title":"Different copper sources\nand levels affect growth performance, copper content, carcass characteristics,\nintestinal microorganism and metabolism of finishing pigs","volume":"8","author":"Wen","year":"2022","journal-title":"Anim. Nutr."},{"key":"2026090104482325300_cit9","doi-asserted-by":"publisher","first-page":"2675","DOI":"10.1007\/s12011-024-04345-8","article-title":"Copper toxicity in animals: a review","volume":"203","author":"Wang","year":"2025","journal-title":"Biol.\nTrace Elem. Res."},{"key":"2026090104482325300_cit10","doi-asserted-by":"publisher","first-page":"353","DOI":"10.1186\/s12964-024-01726-3","article-title":"The crosstalk between\ncopper-induced oxidative stress and cuproptosis: a novel potential\nanticancer paradigm","volume":"22","author":"Vo","year":"2024","journal-title":"Cell Commun. Signal."},{"key":"2026090104482325300_cit11","doi-asserted-by":"publisher","first-page":"981","DOI":"10.3390\/antiox12050981","article-title":"Unfolding the interactions between endoplasmic reticulum\nstress and oxidative stress","volume":"12","author":"Ong","year":"2023","journal-title":"Antioxidants"},{"key":"2026090104482325300_cit12","doi-asserted-by":"publisher","first-page":"123349","DOI":"10.1016\/j.jhazmat.2020.123349","article-title":"Exposure to\ncopper oxide nanoparticles triggers oxidative stress and endoplasmic\nreticulum (ER)-stress induced toxicology and apoptosis in male rat\nliver and BRL-3A cell","volume":"401","author":"Liu","year":"2021","journal-title":"J. Hazard. Mater."},{"key":"2026090104482325300_cit13","doi-asserted-by":"publisher","first-page":"1197","DOI":"10.1007\/s12011-022-03224-4","article-title":"Long-term copper exposure induces\nmitochondrial dynamics disorder and mitophagy in the cerebrum of pigs","volume":"201","author":"Li","year":"2023","journal-title":"Biol. Trace Elem. Res."},{"key":"2026090104482325300_cit14","doi-asserted-by":"publisher","first-page":"3691","DOI":"10.1016\/j.molcel.2021.08.018","article-title":"Redox metabolism: ROS as specific molecular regulators\nof cell signaling and function","volume":"81","author":"Lennicke","year":"2021","journal-title":"Mol. Cell"},{"key":"2026090104482325300_cit15","doi-asserted-by":"publisher","first-page":"14066","DOI":"10.3390\/ijms241814066","article-title":"Unfolded\nprotein response signaling\nin liver disorders: a 2023 updated review","volume":"24","author":"Shreya","year":"2023","journal-title":"Int.\nJ. Mol. Sci."},{"key":"2026090104482325300_cit16","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2412246122","article-title":"Stress-induced\ntranslocation of the endoplasmic reticulum chaperone GRP78\/BiP and\nits impact on human disease and therapy","volume":"122","author":"Lee","year":"2025","journal-title":"Proc.\nNatl. Acad. Sci. U. S. A."},{"key":"2026090104482325300_cit17","doi-asserted-by":"publisher","first-page":"107151","DOI":"10.1016\/j.jbc.2024.107151","article-title":"The integrated\nstress response in metabolic adaptation","volume":"300","author":"Ryoo","year":"2024","journal-title":"J.\nBiol. Chem."},{"key":"2026090104482325300_cit18","doi-asserted-by":"publisher","first-page":"685895","DOI":"10.3389\/fphar.2021.685895","article-title":"CHOP regulates endoplasmic reticulum\nstress-mediated hepatoxicity induced by monocrotaline","volume":"12","author":"Guo","year":"2021","journal-title":"Front. 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Pollut."},{"key":"2026090104482325300_cit23","doi-asserted-by":"publisher","first-page":"1492","DOI":"10.1016\/j.molcel.2022.02.018","article-title":"ER remodeling via ER-phagy","volume":"82","author":"Gubas","year":"2022","journal-title":"Mol. Cell"},{"key":"2026090104482325300_cit24","doi-asserted-by":"publisher","first-page":"576","DOI":"10.1016\/j.tcb.2025.01.002","article-title":"Endoplasmic\nreticulum (ER) protein\ndegradation by ER-associated degradation and ER-phagy","volume":"35","author":"Wu","year":"2025","journal-title":"Trends Cell Biol."},{"key":"2026090104482325300_cit25","doi-asserted-by":"publisher","first-page":"102084","DOI":"10.1016\/j.ceb.2022.102084","article-title":"Regulatory events controlling ER-phagy","volume":"76","author":"Iavarone","year":"2022","journal-title":"Curr. Opin. Cell Biol."},{"key":"2026090104482325300_cit26","doi-asserted-by":"publisher","first-page":"1652","DOI":"10.1038\/s41401-024-01471-w","article-title":"SEC62-dependent ER-phagy contributes\nto apelin-13\/APJ-induced\nmonocyte-vascular endothelial cell adhesion in atherosclerosis pathogenesis","volume":"46","author":"Chen","year":"2025","journal-title":"Acta Pharmacol. Sin."},{"key":"2026090104482325300_cit27","doi-asserted-by":"publisher","first-page":"684526","DOI":"10.3389\/fcell.2021.684526","article-title":"ER-Phagy: a new regulator of ER homeostasis","volume":"9","author":"Yang","year":"2021","journal-title":"Front. Cell. Dev. Biol."},{"key":"2026090104482325300_cit28","doi-asserted-by":"publisher","first-page":"2749","DOI":"10.3390\/ijms24032749","article-title":"The activation\nof reticulophagy by ER stress through the ATF4-MAP1LC3A-CCPG1 pathway\nin ovarian granulosa cells is linked to apoptosis and necroptosis","volume":"24","author":"Li","year":"2023","journal-title":"Int. J. Mol. Sci."},{"key":"2026090104482325300_cit29","doi-asserted-by":"publisher","first-page":"16506","DOI":"10.1021\/acs.jafc.4c03978","article-title":"DRP1 aggravates\ncopper nanoparticle-induced ER-phagy by disturbing mitochondria-associated\nmembranes in chicken hepatocytes","volume":"72","author":"Li","year":"2024","journal-title":"J. Agric.\nFood Chem."},{"key":"2026090104482325300_cit30","doi-asserted-by":"publisher","first-page":"104111","DOI":"10.1016\/j.etap.2023.104111","article-title":"Sodium arsenite-induced\ncytotoxicity is regulated by BNIP3L\/Nix-mediated\nendoplasmic reticulum stress responses and CCPG1-mediated endoplasmic\nreticulum-phagy","volume":"99","author":"Kim","year":"2023","journal-title":"Environ. Toxicol. Pharmacol."},{"key":"2026090104482325300_cit31","doi-asserted-by":"publisher","first-page":"36","DOI":"10.1016\/j.toxlet.2022.05.010","article-title":"Mechanism of ER stress-mediated ER-phagy by CdTe-QDs in yeast cells","volume":"365","author":"Wei","year":"2022","journal-title":"Toxicol. Lett."},{"key":"2026090104482325300_cit32","doi-asserted-by":"publisher","first-page":"1477","DOI":"10.1016\/j.molcel.2022.03.025","article-title":"Reshaping endoplasmic reticulum quality\ncontrol through the unfolded protein response","volume":"82","author":"Wiseman","year":"2022","journal-title":"Mol. Cell"},{"key":"2026090104482325300_cit33","doi-asserted-by":"publisher","first-page":"123713","DOI":"10.1016\/j.envpol.2024.123713","article-title":"Polystyrene nanoplastics and cadmium\nco-exposure aggravated cardiomyocyte damage in mice by regulating\nPANoptosis pathway","volume":"347","author":"Ye","year":"2024","journal-title":"Environ. Pollut."},{"key":"2026090104482325300_cit34","doi-asserted-by":"publisher","first-page":"112040","DOI":"10.1016\/j.ecoenv.2021.112040","article-title":"Metabolomics analysis reveals the effect of copper on autophagy in\nmyocardia of pigs","volume":"213","author":"Li","year":"2021","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"2026090104482325300_cit35","doi-asserted-by":"publisher","first-page":"109710","DOI":"10.1016\/j.ecoenv.2019.109710","article-title":"Effects of copper on oxidative stress\nand autophagy in hypothalamus\nof broilers","volume":"185","author":"Liao","year":"2019","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"2026090104482325300_cit36","doi-asserted-by":"publisher","first-page":"110158","DOI":"10.1016\/j.ecoenv.2019.110158","article-title":"Long-term exposure\nto copper induces autophagy and apoptosis through oxidative stress\nin rat kidneys","volume":"190","author":"Wan","year":"2020","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"2026090104482325300_cit37","doi-asserted-by":"publisher","first-page":"121947","DOI":"10.1016\/j.envpol.2023.121947","article-title":"Co-exposure to environmentally relevant\nconcentrations of cadmium and polystyrene nanoplastics induced oxidative\nstress, ferroptosis and excessive mitophagy in mice kidney","volume":"333","author":"Qiu","year":"2023","journal-title":"Environ. Pollut."},{"key":"2026090104482325300_cit38","doi-asserted-by":"publisher","first-page":"113468","DOI":"10.1016\/j.fct.2022.113468","article-title":"Asiatic acid alleviates\nLPS-induced acute kidney injury in broilers by inhibiting oxidative\nstress and ferroptosis via activation of the Nrf2 pathway","volume":"170","author":"Qiu","year":"2022","journal-title":"Food Chem. Toxicol."},{"key":"2026090104482325300_cit39","doi-asserted-by":"publisher","first-page":"6943","DOI":"10.1021\/acs.jafc.2c01367","article-title":"Latent benefits and\ntoxicity risks transmission chain of high dietary\ncopper along the livestock-environment-plant-human health axis and\nmicrobial homeostasis: a review","volume":"70","author":"Zhen","year":"2022","journal-title":"J. Agric. Food\nChem."},{"key":"2026090104482325300_cit40","doi-asserted-by":"publisher","first-page":"101113","DOI":"10.1016\/j.animal.2024.101113","article-title":"Influence of copper source and dietary inclusion level\non growth performance of weaned pigs and expression of trace element\nrelated genes in the small intestine","volume":"18","author":"Van Baal","year":"2024","journal-title":"Animal"},{"key":"2026090104482325300_cit41","doi-asserted-by":"publisher","first-page":"883","DOI":"10.2147\/JIR.S498340","article-title":"Cuproptosis cell death molecular\nevents and pathways to liver disease","volume":"18","author":"Mao","year":"2025","journal-title":"J. Inflamm.\nRes."},{"key":"2026090104482325300_cit42","doi-asserted-by":"publisher","DOI":"10.1002\/jssc.70312","article-title":"Cell metabolomics reveals\nthe hepatotoxic mechanism\nof copper in normal rat liver cells using reversed-phase and hydrophilic\ninteraction liquid chromatography-quadrupole-time-of-flight mass spectrometry","volume":"48","author":"Liu","year":"2025","journal-title":"J. Sep. Sci."},{"key":"2026090104482325300_cit43","doi-asserted-by":"publisher","first-page":"1633711","DOI":"10.3389\/fimmu.2025.1633711","article-title":"Mechanisms of copper\nmetabolism and cuproptosis: implications for liver diseases","volume":"16","author":"Chen","year":"2025","journal-title":"Front. Immunol."},{"key":"2026090104482325300_cit44","doi-asserted-by":"publisher","first-page":"123688","DOI":"10.1016\/j.lfs.2025.123688","article-title":"Metal ions-induced\nprogrammed cell death: how does\noxidative stress regulate cell death?","volume":"374","author":"Wang","year":"2025","journal-title":"Life\nSci."},{"key":"2026090104482325300_cit45","doi-asserted-by":"publisher","first-page":"114391","DOI":"10.1016\/j.intimp.2025.114391","article-title":"Copper\u2019s\ndual role: reviewing its impact on liver health and disease","volume":"152","author":"Pan","year":"2025","journal-title":"Int. Immunopharmacol."},{"key":"2026090104482325300_cit46","doi-asserted-by":"publisher","first-page":"153","DOI":"10.1007\/s00204-024-03903-2","article-title":"Heavy metals:\ntoxicity and human health effects","volume":"99","author":"Jomova","year":"2025","journal-title":"Arch. 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Res."},{"key":"2026090104482325300_cit49","doi-asserted-by":"publisher","first-page":"111935","DOI":"10.1016\/j.microc.2024.111935","article-title":"Biomarker\nfor the toxicity evaluation of cadmium based on glutathione: recent\nprogress and challenges","volume":"207","author":"Iddrisu","year":"2024","journal-title":"Microchem. J."},{"key":"2026090104482325300_cit50","doi-asserted-by":"publisher","first-page":"563","DOI":"10.1007\/s00204-024-03902-3","article-title":"Therapeutic\npotential\nof 4-phenylbutyric acid against methylmercury-induced neuronal cell\ndeath in mice","volume":"99","author":"Miki","year":"2025","journal-title":"Arch. 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