{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,27]],"date-time":"2026-10-27T03:59:04Z","timestamp":1793073544335,"version":"4.5.0"},"reference-count":60,"publisher":"Springer Science and Business Media LLC","issue":"4","license":[{"start":{"date-parts":[[2026,2,19]],"date-time":"2026-02-19T00:00:00Z","timestamp":1771459200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"},{"start":{"date-parts":[[2026,2,19]],"date-time":"2026-02-19T00:00:00Z","timestamp":1771459200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"}],"funder":[{"DOI":"10.13039\/501100005047","name":"Natural Science Foundation of Liaoning Province","doi-asserted-by":"publisher","award":["201602817"],"award-info":[{"award-number":["201602817"]}],"id":[{"id":"10.13039\/501100005047","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["J Gastroenterol"],"published-print":{"date-parts":[[2026,4]]},"DOI":"10.1007\/s00535-026-02363-y","type":"journal-article","created":{"date-parts":[[2026,2,19]],"date-time":"2026-02-19T13:03:58Z","timestamp":1771506238000},"page":"369-385","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":5,"title":["Targeting the ZDHHC9-mediated STAT1 palmitoylation-phosphorylation conversion inhibits gastric cancer progression"],"prefix":"10.1007","volume":"61","author":[{"given":"Xiao","family":"Bai","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yun-Ran","family":"Guo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wan-Chuan","family":"Zhang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhao-Hu","family":"Shi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5611-7457","authenticated-orcid":false,"given":"Dong-Qiu","family":"Dai","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,2,19]]},"reference":[{"issue":"3","key":"2363_CR1","doi-asserted-by":"publisher","first-page":"209","DOI":"10.3322\/caac.21660","volume":"71","author":"H Sung","year":"2021","unstructured":"Sung H, Ferlay J, Siegel RL, et al. Global cancer statistics 2020: GLOBOCAN estimates of incidence and mortality worldwide for 36 cancers in 185 countries. CA: cancer J Clin. 2021;71(3):209\u201349. https:\/\/doi.org\/10.3322\/caac.21660.","journal-title":"CA: cancer J Clin"},{"key":"2363_CR2","doi-asserted-by":"publisher","first-page":"335","DOI":"10.1053\/j.gastro.2020.02.068","volume":"159","author":"M Arnold","year":"2020","unstructured":"Arnold M, Abnet CC, Neale RE, et al. Global burden of 5 major types of gastrointestinal cancer. Gastroenterology. 2020;159:335-49.e15. https:\/\/doi.org\/10.1053\/j.gastro.2020.02.068.","journal-title":"Gastroenterology"},{"key":"2363_CR3","doi-asserted-by":"publisher","first-page":"105322","DOI":"10.1016\/j.esmoop.2025.105322","volume":"10","author":"S Sato","year":"2025","unstructured":"Sato S, Rhodes SL, Aoki Y, et al. Clinical characterization of FGFR2b expression in patients with advanced gastric or gastroesophageal junction adenocarcinoma. ESMO Open. 2025;10:105322. https:\/\/doi.org\/10.1016\/j.esmoop.2025.105322.","journal-title":"ESMO Open"},{"key":"2363_CR4","doi-asserted-by":"publisher","first-page":"193","DOI":"10.1186\/s12943-025-02397-5","volume":"24","author":"H Jia","year":"2025","unstructured":"Jia H, Jiang L, Shen X, et al. Post-translational modifications of cancer immune checkpoints: mechanisms and therapeutic strategies. Mol Cancer. 2025;24:193. https:\/\/doi.org\/10.1186\/s12943-025-02397-5.","journal-title":"Mol Cancer"},{"issue":"108","key":"2363_CR5","doi-asserted-by":"publisher","first-page":"eado3485","DOI":"10.1126\/sciimmunol.ado3485","volume":"10","author":"J Liang","year":"2025","unstructured":"Liang J, Liao J, Chang R, et al. Riplet promotes lipid metabolism changes associated with CD8 T cell exhaustion and anti-PD-1 resistance in hepatocellular carcinoma. Sci Immunol. 2025;10(108):eado3485. https:\/\/doi.org\/10.1126\/sciimmunol.ado3485.","journal-title":"Sci Immunol"},{"key":"2363_CR6","doi-asserted-by":"publisher","first-page":"4254","DOI":"10.1038\/s41467-025-58552-5","volume":"16","author":"Y Yan","year":"2025","unstructured":"Yan Y, Yu J, Wang W, et al. Palmitoylation prevents B7-H4 lysosomal degradation sustaining tumor immune evasion. Nat Commun. 2025;16:4254. https:\/\/doi.org\/10.1038\/s41467-025-58552-5.","journal-title":"Nat Commun"},{"key":"2363_CR7","doi-asserted-by":"publisher","first-page":"411","DOI":"10.1038\/s42003-020-01145-3","volume":"3","author":"F Plain","year":"2020","unstructured":"Plain F, Howie J, Kennedy J, et al. Control of protein palmitoylation by regulating substrate recruitment to a zDHHC-protein acyltransferase. Commun Biol. 2020;3:411. https:\/\/doi.org\/10.1038\/s42003-020-01145-3.","journal-title":"Commun Biol"},{"key":"2363_CR8","doi-asserted-by":"publisher","first-page":"768","DOI":"10.1038\/s43018-025-00937-y","volume":"6","author":"L Zhou","year":"2025","unstructured":"Zhou L, Lian G, Zhou T, et al. Palmitoylation of GPX4 via the targetable ZDHHC8 determines ferroptosis sensitivity and antitumor immunity. Nat Cancer. 2025;6:768\u201385. https:\/\/doi.org\/10.1038\/s43018-025-00937-y.","journal-title":"Nat Cancer"},{"key":"2363_CR9","doi-asserted-by":"publisher","first-page":"6880","DOI":"10.1158\/0008-5472.Can-17-1536","volume":"77","author":"C Sharma","year":"2017","unstructured":"Sharma C, Wang HX, Li Q, et al. Protein acyltransferase DHHC3 regulates breast tumor growth, oxidative stress, and senescence. Cancer Res. 2017;77:6880\u201390. https:\/\/doi.org\/10.1158\/0008-5472.Can-17-1536.","journal-title":"Cancer Res"},{"key":"2363_CR10","doi-asserted-by":"publisher","first-page":"2126","DOI":"10.1038\/s41388-023-02718-2","volume":"42","author":"Z Lin","year":"2023","unstructured":"Lin Z, Agarwal S, Tan S, et al. Palmitoyl acyltransferase ZDHHC7 inhibits androgen receptor and suppresses prostate cancer. Oncogene. 2023;42:2126\u201338. https:\/\/doi.org\/10.1038\/s41388-023-02718-2.","journal-title":"Oncogene"},{"key":"2363_CR11","doi-asserted-by":"publisher","first-page":"198","DOI":"10.1186\/s13045-021-01214-y","volume":"14","author":"N Awasthi","year":"2021","unstructured":"Awasthi N, Liongue C, Ward AC. STAT proteins: a kaleidoscope of canonical and non-canonical functions in immunity and cancer. J Hematol Oncol. 2021;14:198. https:\/\/doi.org\/10.1186\/s13045-021-01214-y.","journal-title":"J Hematol Oncol"},{"key":"2363_CR12","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1016\/j.semcancer.2019.10.002","volume":"60","author":"Y Verhoeven","year":"2020","unstructured":"Verhoeven Y, Tilborghs S, Jacobs J, et al. The potential and controversy of targeting STAT family members in cancer. Semin Cancer Biol. 2020;60:41\u201356. https:\/\/doi.org\/10.1016\/j.semcancer.2019.10.002.","journal-title":"Semin Cancer Biol"},{"issue":"814","key":"2363_CR13","doi-asserted-by":"publisher","first-page":"eadd2282","DOI":"10.1126\/scisignal.add2282","volume":"16","author":"Y Jiang","year":"2023","unstructured":"Jiang Y, Xu Y, Zhu C, et al. STAT3 palmitoylation initiates a positive feedback loop that promotes the malignancy of hepatocellular carcinoma cells in mice. Sci Signal. 2023;16(814):eadd2282. https:\/\/doi.org\/10.1126\/scisignal.add2282.","journal-title":"Sci Signal"},{"key":"2363_CR14","doi-asserted-by":"publisher","first-page":"6602","DOI":"10.1038\/s41467-025-61875-y","volume":"16","author":"Q Zhang","year":"2025","unstructured":"Zhang Q, Li J, Liu X, et al. Inhibiting CD36 palmitoylation improves cardiac function post-infarction by regulating lipid metabolic homeostasis and autophagy. Nat Commun. 2025;16:6602. https:\/\/doi.org\/10.1038\/s41467-025-61875-y.","journal-title":"Nat Commun"},{"key":"2363_CR15","doi-asserted-by":"publisher","first-page":"113961","DOI":"10.1016\/j.jconrel.2025.113961","volume":"384","author":"X Shi","year":"2025","unstructured":"Shi X, Zhao X, He Y, et al. Posttranslational remodeling micelle reverses cell-surface and exosomal PD-L1 immunosuppression in tumors resistant to PD-L1 antibody therapy. J Control Release. 2025;384:113961. https:\/\/doi.org\/10.1016\/j.jconrel.2025.113961.","journal-title":"J Control Release"},{"key":"2363_CR16","doi-asserted-by":"publisher","first-page":"102208","DOI":"10.1016\/j.xcrm.2025.102208","volume":"6","author":"M Xu","year":"2025","unstructured":"Xu M, Zheng Y, Chen J, et al. CLDN4 palmitoylation promotes hepatic-to-biliary lineage transition and lenvatinib resistance in hepatocellular carcinoma. Cell Rep Med. 2025;6:102208. https:\/\/doi.org\/10.1016\/j.xcrm.2025.102208.","journal-title":"Cell Rep Med"},{"key":"2363_CR17","doi-asserted-by":"publisher","first-page":"717","DOI":"10.1002\/minf.201200150","volume":"32","author":"AA Kaczor","year":"2013","unstructured":"Kaczor AA, Selent J, Sanz F, et al. Modeling complexes of transmembrane proteins: systematic analysis of protein protein docking tools. Mol Inform. 2013;32:717\u201333. https:\/\/doi.org\/10.1002\/minf.201200150.","journal-title":"Mol Inform"},{"key":"2363_CR18","doi-asserted-by":"publisher","first-page":"102463","DOI":"10.1016\/j.sbi.2022.102463","volume":"77","author":"R Puthenveetil","year":"2022","unstructured":"Puthenveetil R, G\u00f3mez-Navarro N, Banerjee A. Access and utilization of long chain fatty acyl-CoA by zDHHC protein acyltransferases. Curr Opin Struct Biol. 2022;77:102463. https:\/\/doi.org\/10.1016\/j.sbi.2022.102463.","journal-title":"Curr Opin Struct Biol"},{"issue":"22","key":"2363_CR19","doi-asserted-by":"publisher","first-page":"jcs251819","DOI":"10.1242\/jcs.251819","volume":"133","author":"C Salaun","year":"2020","unstructured":"Salaun C, Locatelli C, Zmuda F, et al. Accessory proteins of the zDHHC family of S-acylation enzymes. J Cell Sci. 2020;133(22):jcs251819. https:\/\/doi.org\/10.1242\/jcs.251819.","journal-title":"J Cell Sci"},{"key":"2363_CR20","doi-asserted-by":"publisher","first-page":"14640","DOI":"10.1074\/jbc.REV120.014717","volume":"295","author":"F Zmuda","year":"2020","unstructured":"Zmuda F, Chamberlain LH. Regulatory effects of post-translational modifications on zDHHC S-acyltransferases. J Biol Chem. 2020;295:14640\u201352. https:\/\/doi.org\/10.1074\/jbc.REV120.014717.","journal-title":"J Biol Chem"},{"key":"2363_CR21","doi-asserted-by":"publisher","first-page":"203","DOI":"10.1016\/0005-2760(92)90046-x","volume":"1125","author":"RA Coleman","year":"1992","unstructured":"Coleman RA, Rao P, Fogelsong RJ, et al. 2-bromopalmitoyl-CoA and 2-bromopalmitate: promiscuous inhibitors of membrane-bound enzymes. Biochimica et Biophysica Acta (BBA)\u2014Lipids Lipid Metab. 1992;1125:203\u20139. https:\/\/doi.org\/10.1016\/0005-2760(92)90046-x.","journal-title":"Biochimica et Biophysica Acta (BBA)\u2014Lipids Lipid Metab"},{"key":"2363_CR22","doi-asserted-by":"publisher","first-page":"118","DOI":"10.1016\/j.cbpa.2021.07.002","volume":"65","author":"T Lan","year":"2021","unstructured":"Lan T, Delalande C, Dickinson BC. Inhibitors of DHHC family proteins. Curr Opin Chem Biol. 2021;65:118\u201325. https:\/\/doi.org\/10.1016\/j.cbpa.2021.07.002.","journal-title":"Curr Opin Chem Biol"},{"key":"2363_CR23","doi-asserted-by":"publisher","DOI":"10.1016\/j.envpol.2022.120943","author":"Y Zeng","year":"2023","unstructured":"Zeng Y, Chen HQ, Zhang Z, et al. IFI44L as a novel epigenetic silencing tumor suppressor promotes apoptosis through JAK\/STAT1 pathway during lung carcinogenesis. Environ Pollut (Barking, Essex\u202f: 1987). 2023. https:\/\/doi.org\/10.1016\/j.envpol.2022.120943.","journal-title":"Environ Pollut (Barking, Essex : 1987)"},{"key":"2363_CR24","doi-asserted-by":"publisher","first-page":"12373","DOI":"10.1021\/acs.jmedchem.3c00863","volume":"66","author":"R Wang","year":"2023","unstructured":"Wang R, Du TT, Liu WQ, et al. Discovery, optimization, and evaluation of novel N-(Benzimidazol-5-yl)-1,3,4-thiadiazol-2-amine analogues as potent STAT3 inhibitors for cancer treatment. J Med Chem. 2023;66:12373\u201395. https:\/\/doi.org\/10.1021\/acs.jmedchem.3c00863.","journal-title":"J Med Chem"},{"key":"2363_CR25","doi-asserted-by":"publisher","first-page":"12859","DOI":"10.1073\/pnas.192264999","volume":"99","author":"K Ramsauer","year":"2002","unstructured":"Ramsauer K, Sadzak I, Porras A, et al. p38 MAPK enhances STAT1-dependent transcription independently of Ser-727 phosphorylation. Proc Natl Acad Sci U S A. 2002;99:12859\u201364. https:\/\/doi.org\/10.1073\/pnas.192264999.","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"94","key":"2363_CR26","doi-asserted-by":"publisher","first-page":"eadn1452","DOI":"10.1126\/sciimmunol.adn1452","volume":"9","author":"A Balasubramanian","year":"2024","unstructured":"Balasubramanian A, Hsu AY, Ghimire L, et al. The palmitoylation of gasdermin D directs its membrane translocation and pore formation during pyroptosis. Sci Immunol. 2024;9(94):eadn1452. https:\/\/doi.org\/10.1126\/sciimmunol.adn1452.","journal-title":"Sci Immunol"},{"key":"2363_CR27","doi-asserted-by":"publisher","first-page":"757","DOI":"10.1038\/s41556-024-01397-9","volume":"26","author":"N Zhang","year":"2024","unstructured":"Zhang N, Zhang J, Yang Y, et al. A palmitoylation-depalmitoylation relay spatiotemporally controls GSDMD activation in pyroptosis. Nat Cell Biol. 2024;26:757\u201369. https:\/\/doi.org\/10.1038\/s41556-024-01397-9.","journal-title":"Nat Cell Biol"},{"key":"2363_CR28","doi-asserted-by":"publisher","first-page":"545","DOI":"10.1016\/j.tcb.2008.08.008","volume":"18","author":"WX Li","year":"2008","unstructured":"Li WX. Canonical and non-canonical JAK-STAT signaling. Trends Cell Biol. 2008;18:545\u201351. https:\/\/doi.org\/10.1016\/j.tcb.2008.08.008.","journal-title":"Trends Cell Biol"},{"key":"2363_CR29","doi-asserted-by":"publisher","first-page":"602","DOI":"10.1038\/nri1885","volume":"6","author":"NC Reich","year":"2006","unstructured":"Reich NC, Liu L. Tracking STAT nuclear traffic. Nat Rev Immunol. 2006;6:602\u201312. https:\/\/doi.org\/10.1038\/nri1885.","journal-title":"Nat Rev Immunol"},{"key":"2363_CR30","doi-asserted-by":"publisher","first-page":"1314","DOI":"10.1038\/s41467-024-45712-2","volume":"15","author":"Y Zhang","year":"2024","unstructured":"Zhang Y, Wang M, Ye L, et al. HKDC1 promotes tumor immune evasion in hepatocellular carcinoma by coupling cytoskeleton to STAT1 activation and PD-L1 expression. Nat Commun. 2024;15:1314. https:\/\/doi.org\/10.1038\/s41467-024-45712-2.","journal-title":"Nat Commun"},{"key":"2363_CR31","doi-asserted-by":"publisher","first-page":"217475","DOI":"10.1016\/j.canlet.2025.217475","volume":"612","author":"R Wei","year":"2025","unstructured":"Wei R, Song J, Liu C, et al. FAP upregulates PD-L1 expression in cancer-associated fibroblasts to exacerbate T cells dysfunction and suppress anti-tumor immunity. Cancer Lett. 2025;612:217475. https:\/\/doi.org\/10.1016\/j.canlet.2025.217475.","journal-title":"Cancer Lett"},{"key":"2363_CR32","doi-asserted-by":"publisher","first-page":"2087","DOI":"10.1016\/s0140-6736(25)00052-2","volume":"405","author":"R Sundar","year":"2025","unstructured":"Sundar R, Nakayama I, Markar SR, et al. Gastric cancer. Lancet. 2025;405:2087\u2013102. https:\/\/doi.org\/10.1016\/s0140-6736(25)00052-2.","journal-title":"Lancet"},{"issue":"5","key":"2363_CR33","doi-asserted-by":"publisher","first-page":"169","DOI":"10.6004\/jnccn.2025.0022","volume":"23","author":"JA Ajani","year":"2025","unstructured":"Ajani JA, D\u2019Amico TA, Bentrem DJ, et al. Gastric cancer, version 2.2025, NCCN clinical practice guidelines in oncology. J Natl Compr Cancer Netw: JNCCN. 2025;23(5):169\u201391. https:\/\/doi.org\/10.6004\/jnccn.2025.0022.","journal-title":"J Natl Compr Cancer Netw: JNCCN"},{"issue":"101","key":"2363_CR34","doi-asserted-by":"publisher","first-page":"eadp7302","DOI":"10.1126\/sciimmunol.adp7302","volume":"9","author":"Z Zhang","year":"2024","unstructured":"Zhang Z, Ren C, Xiao R, et al. Palmitoylation of TIM-3 promotes immune exhaustion and restrains antitumor immunity. Sci Immunol. 2024;9(101):eadp7302. https:\/\/doi.org\/10.1126\/sciimmunol.adp7302.","journal-title":"Sci Immunol"},{"key":"2363_CR35","doi-asserted-by":"publisher","first-page":"2034","DOI":"10.1158\/0008-5472.Can-22-3105","volume":"83","author":"Y Sun","year":"2023","unstructured":"Sun Y, Zhu L, Liu P, et al. ZDHHC2-mediated AGK palmitoylation activates AKT-mTOR signaling to reduce sunitinib sensitivity in renal cell carcinoma. Cancer Res. 2023;83:2034\u201351. https:\/\/doi.org\/10.1158\/0008-5472.Can-22-3105.","journal-title":"Cancer Res"},{"key":"2363_CR36","doi-asserted-by":"publisher","first-page":"1156","DOI":"10.1136\/gutjnl-2023-330826","volume":"73","author":"L Bu","year":"2024","unstructured":"Bu L, Zhang Z, Chen J, et al. High-fat diet promotes liver tumorigenesis via palmitoylation and activation of AKT. Gut. 2024;73:1156\u201368. https:\/\/doi.org\/10.1136\/gutjnl-2023-330826.","journal-title":"Gut"},{"key":"2363_CR37","doi-asserted-by":"publisher","DOI":"10.1016\/j.metabol.2024.155905","author":"Y Yang","year":"2024","unstructured":"Yang Y, Liu X, Yang D, et al. Interplay of CD36, autophagy, and lipid metabolism: insights into cancer progression. Metab Clin Exp. 2024. https:\/\/doi.org\/10.1016\/j.metabol.2024.155905.","journal-title":"Metab Clin Exp"},{"key":"2363_CR38","doi-asserted-by":"publisher","first-page":"103051","DOI":"10.1016\/j.redox.2024.103051","volume":"70","author":"L Liu","year":"2024","unstructured":"Liu L, Wang L, Liu L, et al. Acyltransferase zinc finger DHHC-type containing 2 aggravates gastric carcinoma growth by targeting Nrf2 signaling: a mechanism-based multicombination bionic nano-drug therapy. Redox Biol. 2024;70:103051. https:\/\/doi.org\/10.1016\/j.redox.2024.103051.","journal-title":"Redox Biol"},{"key":"2363_CR39","doi-asserted-by":"publisher","first-page":"403","DOI":"10.3892\/or.2014.3166","volume":"32","author":"HZ Oo","year":"2014","unstructured":"Oo HZ, Sentani K, Sakamoto N, et al. Overexpression of ZDHHC14 promotes migration and invasion of scirrhous type gastric cancer. Oncol Rep. 2014;32:403\u201310. https:\/\/doi.org\/10.3892\/or.2014.3166.","journal-title":"Oncol Rep"},{"key":"2363_CR40","doi-asserted-by":"publisher","first-page":"477","DOI":"10.1038\/s42003-025-07897-0","volume":"8","author":"H Guo","year":"2025","unstructured":"Guo H, Lin Z, Zhang D, et al. ZDHHC9-mediated CD38 palmitoylation stabilizes CD38 expression and promotes pancreatic cancer growth. Commun Biol. 2025;8:477. https:\/\/doi.org\/10.1038\/s42003-025-07897-0.","journal-title":"Commun Biol"},{"key":"2363_CR41","doi-asserted-by":"publisher","first-page":"217118","DOI":"10.1016\/j.canlet.2024.217118","volume":"598","author":"W Li","year":"2024","unstructured":"Li W, Liu J, Yu T, et al. ZDHHC9-mediated Bip\/GRP78 S-palmitoylation inhibits unfolded protein response and promotes bladder cancer progression. Cancer Lett. 2024;598:217118. https:\/\/doi.org\/10.1016\/j.canlet.2024.217118.","journal-title":"Cancer Lett"},{"key":"2363_CR42","doi-asserted-by":"publisher","first-page":"5872","DOI":"10.1038\/s41467-021-26180-4","volume":"12","author":"Z Zhang","year":"2021","unstructured":"Zhang Z, Li X, Yang F, et al. DHHC9-mediated GLUT1 S-palmitoylation promotes glioblastoma glycolysis and tumorigenesis. Nat Commun. 2021;12:5872. https:\/\/doi.org\/10.1038\/s41467-021-26180-4.","journal-title":"Nat Commun"},{"key":"2363_CR43","doi-asserted-by":"publisher","first-page":"216696","DOI":"10.1016\/j.canlet.2024.216696","volume":"587","author":"L Chen","year":"2024","unstructured":"Chen L, Xing X, Zhu Y, et al. Palmitoylation alters LDHA activity and pancreatic cancer response to chemotherapy. Cancer Lett. 2024;587:216696. https:\/\/doi.org\/10.1016\/j.canlet.2024.216696.","journal-title":"Cancer Lett"},{"key":"2363_CR44","doi-asserted-by":"publisher","first-page":"1896","DOI":"10.1038\/sj.bjc.6603818","volume":"96","author":"F Mansilla","year":"2007","unstructured":"Mansilla F, Birkenkamp-Demtroder K, Kruh\u00f8ffer M, et al. Differential expression of DHHC9 in microsatellite stable and instable human colorectal cancer subgroups. Br J Cancer. 2007;96:1896\u2013903. https:\/\/doi.org\/10.1038\/sj.bjc.6603818.","journal-title":"Br J Cancer"},{"key":"2363_CR45","doi-asserted-by":"publisher","first-page":"894","DOI":"10.1093\/bioinformatics\/btl013","volume":"22","author":"F Zhou","year":"2006","unstructured":"Zhou F, Xue Y, Yao X, et al. CSS-Palm: palmitoylation site prediction with a clustering and scoring strategy (CSS). Bioinformatics. 2006;22:894\u20136. https:\/\/doi.org\/10.1093\/bioinformatics\/btl013.","journal-title":"Bioinformatics"},{"key":"2363_CR46","doi-asserted-by":"publisher","first-page":"1189","DOI":"10.1016\/j.celrep.2017.04.031","volume":"19","author":"A Garcia-Diaz","year":"2017","unstructured":"Garcia-Diaz A, Shin DS, Moreno BH, et al. Interferon receptor signaling pathways regulating PD-L1 and PD-L2 expression. Cell Rep. 2017;19:1189\u2013201. https:\/\/doi.org\/10.1016\/j.celrep.2017.04.031.","journal-title":"Cell Rep"},{"key":"2363_CR47","doi-asserted-by":"publisher","first-page":"7","DOI":"10.1007\/s00262-023-03624-y","volume":"73","author":"Y Wen","year":"2024","unstructured":"Wen Y, Ye S, Li Z, et al. HDAC6 inhibitor ACY-1215 enhances STAT1 acetylation to block PD-L1 for colorectal cancer immunotherapy. Cancer Immunol Immunother. 2024;73:7. https:\/\/doi.org\/10.1007\/s00262-023-03624-y.","journal-title":"Cancer Immunol Immunother"},{"key":"2363_CR48","doi-asserted-by":"publisher","first-page":"123","DOI":"10.1002\/cac2.12386","volume":"43","author":"L Zhang","year":"2023","unstructured":"Zhang L, Li Q, Yang J, et al. Cytosolic TGM2 promotes malignant progression in gastric cancer by suppressing the TRIM21-mediated ubiquitination\/degradation of STAT1 in a GTP binding-dependent modality. Cancer Commun. 2023;43:123\u201349. https:\/\/doi.org\/10.1002\/cac2.12386.","journal-title":"Cancer Commun"},{"key":"2363_CR49","doi-asserted-by":"publisher","first-page":"1844","DOI":"10.1016\/j.jaci.2017.07.027","volume":"141","author":"EP Sampaio","year":"2018","unstructured":"Sampaio EP, Ding L, Rose SR, et al. Novel signal transducer and activator of transcription 1 mutation disrupts small ubiquitin-related modifier conjugation causing gain of function. J Allergy Clin Immunol. 2018;141:1844-53.e2. https:\/\/doi.org\/10.1016\/j.jaci.2017.07.027.","journal-title":"J Allergy Clin Immunol"},{"key":"2363_CR50","doi-asserted-by":"publisher","first-page":"207","DOI":"10.1186\/s12943-024-02124-6","volume":"23","author":"X Zang","year":"2024","unstructured":"Zang X, He XY, Xiao CM, et al. Circular RNA-encoded oncogenic PIAS1 variant blocks immunogenic ferroptosis by modulating the balance between SUMOylation and phosphorylation of STAT1. Mol Cancer. 2024;23:207. https:\/\/doi.org\/10.1186\/s12943-024-02124-6.","journal-title":"Mol Cancer"},{"key":"2363_CR51","doi-asserted-by":"publisher","first-page":"40","DOI":"10.1016\/j.mce.2009.10.007","volume":"315","author":"OH Kr\u00e4mer","year":"2010","unstructured":"Kr\u00e4mer OH, Heinzel T. Phosphorylation-acetylation switch in the regulation of STAT1 signaling. Mol Cell Endocrinol. 2010;315:40\u20138. https:\/\/doi.org\/10.1016\/j.mce.2009.10.007.","journal-title":"Mol Cell Endocrinol"},{"key":"2363_CR52","doi-asserted-by":"publisher","first-page":"116367","DOI":"10.1016\/j.bcp.2024.116367","volume":"226","author":"K Wang","year":"2024","unstructured":"Wang K, Chen Y, Zhang M, et al. Metformin suppresses gastric cancer progression by disrupting the STAT1-PRMT1 axis. Biochem Pharmacol. 2024;226:116367. https:\/\/doi.org\/10.1016\/j.bcp.2024.116367.","journal-title":"Biochem Pharmacol"},{"key":"2363_CR53","doi-asserted-by":"publisher","first-page":"25678","DOI":"10.1074\/jbc.M116.756544","volume":"291","author":"W Zheng","year":"2016","unstructured":"Zheng W, Yang J, Beauchamp E, et al. Regulation of TRPP3 channel function by N-terminal domain palmitoylation and phosphorylation. J Biol Chem. 2016;291:25678\u201391. https:\/\/doi.org\/10.1074\/jbc.M116.756544.","journal-title":"J Biol Chem"},{"key":"2363_CR54","doi-asserted-by":"publisher","first-page":"221","DOI":"10.1038\/s41392-024-01934-w","volume":"9","author":"Y Lv","year":"2024","unstructured":"Lv Y, Qi J, Babon JJ, et al. The JAK-STAT pathway: from structural biology to cytokine engineering. Signal Transduct Target Ther. 2024;9:221. https:\/\/doi.org\/10.1038\/s41392-024-01934-w.","journal-title":"Signal Transduct Target Ther"},{"key":"2363_CR55","doi-asserted-by":"publisher","first-page":"31141","DOI":"10.1074\/jbc.M504113200","volume":"280","author":"JT Swarthout","year":"2005","unstructured":"Swarthout JT, Lobo S, Farh L, et al. DHHC9 and GCP16 constitute a human protein fatty acyltransferase with specificity for H and N-Ras. J Biol Chem. 2005;280:31141\u20138. https:\/\/doi.org\/10.1074\/jbc.M504113200.","journal-title":"J Biol Chem"},{"key":"2363_CR56","doi-asserted-by":"publisher","first-page":"83","DOI":"10.1038\/s41422-018-0124-5","volume":"29","author":"Y Yang","year":"2019","unstructured":"Yang Y, Hsu JM, Sun L, et al. Palmitoylation stabilizes PD-L1 to promote breast tumor growth. Cell Res. 2019;29:83\u20136. https:\/\/doi.org\/10.1038\/s41422-018-0124-5.","journal-title":"Cell Res"},{"key":"2363_CR57","doi-asserted-by":"publisher","DOI":"10.1016\/j.biopha.2023.114567","author":"Z Lin","year":"2023","unstructured":"Lin Z, Huang K, Guo H, et al. Targeting ZDHHC9 potentiates anti-programmed death-ligand 1 immunotherapy of pancreatic cancer by modifying the tumor microenvironment. Biomed Pharmacother. 2023. https:\/\/doi.org\/10.1016\/j.biopha.2023.114567.","journal-title":"Biomed Pharmacother"},{"key":"2363_CR58","doi-asserted-by":"publisher","first-page":"306","DOI":"10.1038\/s41551-019-0375-6","volume":"3","author":"H Yao","year":"2019","unstructured":"Yao H, Lan J, Li C, et al. Inhibiting PD-L1 palmitoylation enhances T-cell immune responses against tumours. Nat Biomed Eng. 2019;3:306\u201317. https:\/\/doi.org\/10.1038\/s41551-019-0375-6.","journal-title":"Nat Biomed Eng"},{"key":"2363_CR59","doi-asserted-by":"publisher","DOI":"10.20517\/2394-4722.2020.58","author":"Z Qureshy","year":"2020","unstructured":"Qureshy Z, Johnson DE, Grandis JR. Targeting the JAK\/STAT pathway in solid tumors. J Cancer Metastasis Treat. 2020. https:\/\/doi.org\/10.20517\/2394-4722.2020.58.","journal-title":"J Cancer Metastasis Treat"},{"key":"2363_CR60","doi-asserted-by":"publisher","first-page":"365","DOI":"10.1038\/bjc.2015.233","volume":"113","author":"SJ Thomas","year":"2015","unstructured":"Thomas SJ, Snowden JA, Zeidler MP, et al. The role of JAK\/STAT signalling in the pathogenesis, prognosis and treatment of solid tumours. Br J Cancer. 2015;113:365\u201371. https:\/\/doi.org\/10.1038\/bjc.2015.233.","journal-title":"Br J Cancer"}],"container-title":["Journal of Gastroenterology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00535-026-02363-y.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s00535-026-02363-y","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00535-026-02363-y.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,4,3]],"date-time":"2026-04-03T10:50:57Z","timestamp":1775213457000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s00535-026-02363-y"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,2,19]]},"references-count":60,"journal-issue":{"issue":"4","published-print":{"date-parts":[[2026,4]]}},"alternative-id":["2363"],"URL":"https:\/\/doi.org\/10.1007\/s00535-026-02363-y","relation":{},"ISSN":["0944-1174","1435-5922"],"issn-type":[{"value":"0944-1174","type":"print"},{"value":"1435-5922","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,2,19]]},"assertion":[{"value":"2 September 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"6 February 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"19 February 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare that they have no competing interests.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Conflict of interest"}},{"value":"This study complied with the Declaration of Helsinki and was approved by the Ethics Committee of the Fourth Affiliated Hospital (EC-2025-KS-090). All animal experiments were approved by the Ethics Committee of China Medical University (CMU20241811).","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethical approval and consent to participate"}},{"value":"Not applicable.","order":4,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent for publication"}}]}}