{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,21]],"date-time":"2026-07-21T04:44:47Z","timestamp":1784609087116,"version":"3.55.0"},"reference-count":168,"publisher":"MDPI AG","issue":"2","license":[{"start":{"date-parts":[[2026,1,19]],"date-time":"2026-01-19T00:00:00Z","timestamp":1768780800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"national key specialized project","award":["GJ202302"],"award-info":[{"award-number":["GJ202302"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Keratin, a core structural protein in epithelial cells, is essential for maintaining epithelial tissue integrity. Numerous studies have confirmed its critical role in proliferative disorders, including lung\/liver cancer, idiopathic pulmonary fibrosis (IPF), and hepatic fibrosis (HF). Post-translational modification (PTM) regulates protein activity, and keratin undergoes phosphorylation, acetylation, and methylation\u2014modifications that modulate fibrosis and cancer progression by regulating relevant signaling pathways. However, how these modifications reshape keratin\u2019s structure and function in these diseases remains understudied, underscoring the necessity for a systematic review. This review first summarizes keratin\u2019s classification, physiological functions, and roles in epithelial cells, then focuses on the physiological significance of keratin modifications in fibrosis and cancer, while dissecting the molecular mechanisms by which keratin PTMs drive disease progression to address the knowledge gap regarding modification-related keratin changes. Elucidating the mechanisms of keratin and its PTMs is pivotal for understanding disease progression and developing targeted therapies; meanwhile, keratin-targeted strategies\u2014such as keratin siRNAs and small-molecule compounds that regulate keratin expression or modification\u2014have shown therapeutic potential. In summary, this review synthesizes current research findings and provides novel insights for the treatment of fibrosis and cancer.<\/jats:p>","DOI":"10.3390\/ijms27020972","type":"journal-article","created":{"date-parts":[[2026,1,20]],"date-time":"2026-01-20T11:27:58Z","timestamp":1768908478000},"page":"972","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Regulation of Keratin Chemical Modifications: Potential Molecular Mechanisms in Proliferative Diseases"],"prefix":"10.3390","volume":"27","author":[{"given":"Xuemei","family":"Ma","sequence":"first","affiliation":[{"name":"Department of Basic Medicine, Jiamusi University, Jiamusi 154007, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaoli","family":"Jiang","sequence":"additional","affiliation":[{"name":"Department of Basic Medicine, Jiamusi University, Jiamusi 154007, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mengxue","family":"Song","sequence":"additional","affiliation":[{"name":"Department of Clinical Medicine, Jiamusi University, Jiamusi 154007, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bingbing","family":"Bai","sequence":"additional","affiliation":[{"name":"Department of Basic Medicine, Jiamusi University, Jiamusi 154007, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xia","family":"Hou","sequence":"additional","affiliation":[{"name":"Department of Basic Medicine, Jiamusi University, Jiamusi 154007, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qingtian","family":"Wu","sequence":"additional","affiliation":[{"name":"Department of Basic Medicine, Jiamusi University, Jiamusi 154007, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,1,19]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","unstructured":"Antar, S.A., Ashour, N.A., Marawan, M.E., and Al-Karmalawy, A.A. (2023). Fibrosis: Types, Effects, Markers, Mechanisms for Disease Progression, and Its Relation with Oxidative Stress, Immunity, and Inflammation. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms24044004"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"122745","DOI":"10.1016\/j.lfs.2024.122745","article-title":"The role and molecular mechanism of CTHRC1 in fibrosis","volume":"350","author":"Cao","year":"2024","journal-title":"Life Sci."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.matbio.2018.04.009","article-title":"Decoding fibrosis: Mechanisms and translational aspects","volume":"68\u201369","author":"Schaefer","year":"2018","journal-title":"Matrix Biol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"2","DOI":"10.1016\/j.mam.2018.06.003","article-title":"Organ and tissue fibrosis: Molecular signals, cellular mechanisms and translational implications","volume":"65","author":"Weiskirchen","year":"2019","journal-title":"Mol. Aspects Med."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Dituri, F., Cossu, C., Mancarella, S., and Giannelli, G. (2019). The Interactivity between TGF\u03b2 and BMP Signaling in Organogenesis, Fibrosis, and Cancer. Cells, 8.","DOI":"10.3390\/cells8101130"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"1024","DOI":"10.1093\/ndt\/gfaa301","article-title":"Fibrosis and cancer: Shared features and mechanisms suggest common targeted therapeutic approaches","volume":"37","author":"Landolt","year":"2022","journal-title":"Nephrol. Dial. Transplant."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Yasuma, T., and Gabazza, E.C. (2022). Chronic Fibrosis and Its Progression to Cancer. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms23073924"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"33","DOI":"10.3350\/cmh.2022.0058","article-title":"Wnt signaling in liver regeneration, disease, and cancer","volume":"29","author":"Zou","year":"2023","journal-title":"Clin. Mol. Hepatol."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1038\/s41416-020-01172-1","article-title":"The fibrotic and immune microenvironments as targetable drivers of metastasis","volume":"124","author":"Boulter","year":"2021","journal-title":"Br. J. Cancer"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"555","DOI":"10.1038\/s41586-020-2938-9","article-title":"Fibrosis: From mechanisms to medicines","volume":"587","author":"Henderson","year":"2020","journal-title":"Nature"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1038\/s41573-019-0040-5","article-title":"Targeting metabolic dysregulation for fibrosis therapy","volume":"19","author":"Zhao","year":"2020","journal-title":"Nat. Rev. Drug Discov."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"a018275","DOI":"10.1101\/cshperspect.a018275","article-title":"Types I and II Keratin Intermediate Filaments","volume":"10","author":"Jacob","year":"2018","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"102264","DOI":"10.1016\/j.ceb.2023.102264","article-title":"Posttranslational modifications of keratins and their associated proteins as therapeutic targets in keratin diseases","volume":"85","author":"Li","year":"2023","journal-title":"Curr. Opin. Cell Biol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"2485","DOI":"10.1080\/15548627.2023.2186099","article-title":"Phosphorylation of K8 (keratin 8) by excessive mechanical load-activated PKN (protein kinase N) impairs autophagosome initiation and contributes to disc degeneration","volume":"19","author":"Wang","year":"2023","journal-title":"Autophagy"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"163","DOI":"10.1038\/nrm3753","article-title":"Post-translational modifications of intermediate filament proteins: Mechanisms and functions","volume":"15","author":"Snider","year":"2014","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"102282","DOI":"10.1016\/j.ceb.2023.102282","article-title":"The keratin-desmosome scaffold of internal epithelia in health and disease\u2014The plot is thickening","volume":"86","author":"Toivola","year":"2024","journal-title":"Curr. Opin. Cell Biol."},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Zhang, H., Zhang, Y., Xia, T., Lu, L., Luo, M., Chen, Y., Liu, Y., and Li, Y. (2022). The Role of Keratin17 in Human Tumours. Front. Cell Dev. Biol., 10.","DOI":"10.3389\/fcell.2022.818416"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"110","DOI":"10.1186\/s12964-023-01380-1","article-title":"Crosstalk between protein post-translational modifications and phase separation","volume":"22","author":"Liu","year":"2024","journal-title":"Cell Commun. Signal."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"2148","DOI":"10.1242\/jcs.080127","article-title":"Keratin 8 phosphorylation regulates keratin reorganization and migration of epithelial tumor cells","volume":"125","author":"Busch","year":"2012","journal-title":"J. Cell Sci."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"808","DOI":"10.1097\/HEP.0000000000000875","article-title":"Nuclear K19 is a transcriptional corepressor promoting histone deacetylation and liver tumorigenesis","volume":"81","author":"Han","year":"2025","journal-title":"Hepatology"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"9030","DOI":"10.1096\/fj.201800263RR","article-title":"Liver disease-associated keratin 8 and 18 mutations modulate keratin acetylation and methylation","volume":"33","author":"Jang","year":"2019","journal-title":"FASEB J."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"1065","DOI":"10.1021\/acscentsci.4c00163","article-title":"O-GlcNAcylation Facilitates the Interaction between Keratin 18 and Isocitrate Dehydrogenases and Potentially Influencing Cholangiocarcinoma Progression","volume":"10","author":"Meng","year":"2024","journal-title":"ACS Cent. Sci."},{"key":"ref_23","first-page":"643","article-title":"Keratins are asymmetrically inherited fate determinants in the mammalian embryo","volume":"584","author":"Plachta","year":"2020","journal-title":"Nature"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"105878","DOI":"10.1016\/j.biocel.2020.105878","article-title":"Keratin intermediate filaments in the colon: Guardians of epithelial homeostasis","volume":"129","author":"Polari","year":"2020","journal-title":"Int. J. Biochem. Cell Biol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1186\/s40246-021-00374-9","article-title":"Update of the keratin gene family: Evolution, tissue-specific expression patterns, and relevance to clinical disorders","volume":"16","author":"Ho","year":"2022","journal-title":"Hum. Genom."},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Kalabusheva, E.P., Shtompel, A.S., Rippa, A.L., Ulianov, S.V., Razin, S.V., and Vorotelyak, E.A. (2023). A Kaleidoscope of Keratin Gene Expression and the Mosaic of Its Regulatory Mechanisms. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms24065603"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"56","DOI":"10.1016\/j.ceb.2014.12.006","article-title":"Regulation of keratin network organization","volume":"32","author":"Loschke","year":"2015","journal-title":"Curr. Opin. Cell Biol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"395","DOI":"10.1016\/j.ijbiomac.2022.08.002","article-title":"Biological importance and pharmaceutical significance of keratin: A review","volume":"219","author":"Sarma","year":"2022","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"1103","DOI":"10.1016\/j.molmed.2025.03.012","article-title":"Keratinocytes: New perspectives in inflammatory skin diseases","volume":"31","author":"Ye","year":"2025","journal-title":"Trends Mol. Med."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"6259","DOI":"10.1038\/s41467-024-50481-z","article-title":"Loss-of-function mutations in Keratin 32 gene disrupt skin immune homeostasis in pityriasis rubra pilaris","volume":"15","author":"Shi","year":"2024","journal-title":"Nat. Commun."},{"key":"ref_31","first-page":"Jeb244568","article-title":"The human keratin family: From sequence to function and pathology","volume":"225","author":"Moll","year":"2022","journal-title":"J. Exp. Biol."},{"key":"ref_32","first-page":"583","article-title":"Regulatory networks of keratin genes in epithelial differentiation and disease","volume":"23","author":"Lee","year":"2022","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"102075","DOI":"10.1016\/j.labinv.2024.102075","article-title":"Keratin 6A Is Expressed at the Invasive Front and Enhances the Progression of Colorectal Cancer","volume":"104","author":"Kouchi","year":"2024","journal-title":"Lab. Investig."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"2264","DOI":"10.1016\/j.csbj.2025.05.042","article-title":"Integrated transcriptome and proteome analysis identifies keratins as key regulators of corneal scarring in a murine model","volume":"27","author":"Zhou","year":"2025","journal-title":"Comput. Struct. Biotechnol. J."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"816","DOI":"10.1016\/j.semcancer.2021.12.006","article-title":"Vimentin and cytokeratin: Good alone, bad together","volume":"86","author":"Kuburich","year":"2022","journal-title":"Semin. Cancer Biol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"100817","DOI":"10.1016\/j.mam.2019.09.001","article-title":"Epithelial keratins: Biology and implications as diagnostic markers for liquid biopsies","volume":"72","author":"Werner","year":"2020","journal-title":"Mol. Asp. Med."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"jcs260594","DOI":"10.1242\/jcs.260594","article-title":"Revisiting the significance of keratin expression in complex epithelia","volume":"135","author":"Cohen","year":"2022","journal-title":"J. Cell Sci."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"107607","DOI":"10.1016\/j.biotechadv.2020.107607","article-title":"Microbial enzymes catalyzing keratin degradation: Classification, structure, function","volume":"44","author":"Qiu","year":"2020","journal-title":"Biotechnol. Adv."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"110612","DOI":"10.1016\/j.msec.2019.110612","article-title":"Keratinous materials: Structures and functions in biomedical applications","volume":"110","author":"Rajabi","year":"2020","journal-title":"Mater. Sci. Eng. C Mater. Biol. Appl."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"80","DOI":"10.1016\/j.semcdb.2021.09.018","article-title":"Assembly and recognition of keratins: A structural perspective","volume":"128","author":"Yu","year":"2022","journal-title":"Semin. Cell Dev. Biol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"188664","DOI":"10.1016\/j.bbcan.2021.188664","article-title":"Keratin 1 as a cell-surface receptor in cancer","volume":"1877","author":"Ogunnigbagbe","year":"2022","journal-title":"Biochim. Biophys. Acta (BBA)\u2014Rev. Cancer"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"271","DOI":"10.1016\/j.str.2020.02.002","article-title":"The Interface between Keratin Structurotype and Human Disease","volume":"28","author":"Eldirany","year":"2020","journal-title":"Structure"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"E149","DOI":"10.1111\/jdv.14639","article-title":"Autosomal recessive epidermolysis bullosa simplex due to K14 mutation: Two large Palestinian families and literature review","volume":"32","author":"Diociaiuti","year":"2018","journal-title":"J. Eur. Acad. Dermatol. Venereol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"262","DOI":"10.1111\/bjd.15053","article-title":"Digenic inheritance in epidermolysis bullosa simplex involving two novel mutations in K5 and K14","volume":"177","author":"Kim","year":"2017","journal-title":"Br. J. Dermatol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1146\/annurev-cellbio-100617-062534","article-title":"Cytoplasmic Intermediate Filaments in Cell Biology","volume":"34","year":"2018","journal-title":"Annu. Rev. Cell Dev. Biol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"eabd6187","DOI":"10.1126\/sciadv.abd6187","article-title":"The keratin network of intermediate filaments regulates keratinocyte rigidity sensing and nuclear mechanotransduction","volume":"7","author":"Laly","year":"2021","journal-title":"Sci. Adv."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"2759","DOI":"10.1016\/j.devcel.2024.06.011","article-title":"Shifts in keratin isoform expression activate motility signals during wound healing","volume":"59","author":"Nanes","year":"2024","journal-title":"Dev. Cell"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"102070","DOI":"10.3748\/wjg.v31.i6.102070","article-title":"Keratin 1 modulates intestinal barrier and immune response via kallikrein kinin system in ulcerative colitis","volume":"31","author":"Dong","year":"2025","journal-title":"World J. Gastroenterol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"933","DOI":"10.1038\/ng.3355","article-title":"Keratin-dependent regulation of Aire and gene expression in skin tumor keratinocytes","volume":"47","author":"Hobbs","year":"2015","journal-title":"Nat. Genet."},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Zhang, X., Yin, M., and Zhang, L.-J. (2019). Keratin 6, 16 and 17-Critical Barrier Alarmin Molecules in Skin Wounds and Psoriasis. Cells, 8.","DOI":"10.3390\/cells8080807"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"36","DOI":"10.1007\/s12272-021-01305-x","article-title":"Skin barrier dysfunction and filaggrin","volume":"44","author":"Kim","year":"2021","journal-title":"Arch. Pharmacal Res."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"G438","DOI":"10.1152\/ajpgi.00220.2023","article-title":"Colonocyte keratins stabilize mitochondria and contribute to mitochondrial energy metabolism","volume":"327","author":"Thapa","year":"2024","journal-title":"Am. J. Physiol. Gastrointest. Liver Physiol."},{"key":"ref_53","doi-asserted-by":"crossref","unstructured":"Pang, B., Zhu, Z., Xiao, C., Luo, Y., Fang, H., Bai, Y., Sun, Z., Ma, J., Dang, E., and Wang, G. (2021). Keratin 17 Is Required for Lipid Metabolism in Keratinocytes and Benefits Epidermal Permeability Barrier Homeostasis. Front. Cell Dev. Biol., 9.","DOI":"10.3389\/fcell.2021.779257"},{"key":"ref_54","doi-asserted-by":"crossref","unstructured":"Seiringer, P., Hillig, C., Sch\u00e4bitz, A., Jargosch, M., Pilz, A.C., Eyerich, S., Szegedi, A., Sochorov\u00e1, M., Gruber, F., and Zouboulis, C.C. (2024). Spatial transcriptomics reveals altered lipid metabolism and inflammation-related gene expression of sebaceous glands in psoriasis and atopic dermatitis. Front. Immunol., 15.","DOI":"10.3389\/fimmu.2024.1334844"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"2128","DOI":"10.1080\/15548627.2020.1816342","article-title":"Keratinocyte autophagy enables the activation of keratinocytes and fibroblastsand facilitates wound healing","volume":"17","author":"Qiang","year":"2021","journal-title":"Autophagy"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"257","DOI":"10.1111\/wrr.13137","article-title":"Human keratin matrices promote wound healing by modulating skin cell expression of cytokines and growth factors","volume":"32","author":"Walthall","year":"2024","journal-title":"Wound Repair Regen."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"141752","DOI":"10.1016\/j.ijbiomac.2025.141752","article-title":"Keratin: A potential driver of tumor metastasis","volume":"307","author":"Li","year":"2025","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1186\/s13058-022-01502-6","article-title":"K13 promotes stemness and drives metastasis in breast cancer through a plakoglobin\/c-Myc signaling pathway","volume":"24","author":"Yin","year":"2022","journal-title":"Breast Cancer Res."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"1440","DOI":"10.1164\/rccm.202107-1769OC","article-title":"CYFRA 21-1 Predicts Progression in Idiopathic Pulmonary Fibrosis: A Prospective Longitudinal Analysis of the PROFILE Cohort","volume":"205","author":"Molyneaux","year":"2022","journal-title":"Am. J. Respir. Crit. Care Med."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"120147","DOI":"10.1016\/j.cca.2025.120147","article-title":"Cytokeratin 18 fragment in liver inflammation and fibrosis: Systematic review and meta-analysis","volume":"569","author":"Ye","year":"2025","journal-title":"Clin. Chim. Acta"},{"key":"ref_61","first-page":"637","article-title":"Mutations in K5 and K14 cause epidermolysis bullosa simplex in 75% of the patients","volume":"164","author":"Bolling","year":"2011","journal-title":"Br. J. Dermatol."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"102133","DOI":"10.1016\/j.labinv.2024.102133","article-title":"Impact of Keratins 8 and 18 Genetic Variants on the Severity of Alcoholic Liver Disease","volume":"104","author":"Tihy","year":"2024","journal-title":"Lab. Investig."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1016\/j.semcdb.2021.06.017","article-title":"Keratose hydrogel for tissue regeneration and drug delivery","volume":"128","author":"Ledford","year":"2022","journal-title":"Semin. Cell Dev. Biol."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"3146","DOI":"10.1016\/j.jid.2022.06.022","article-title":"Comparative Spatial Transcriptomic and Single-Cell Analyses of Human Nail Units and Hair Follicles Show Transcriptional Similarities between the Onychodermis and Follicular Dermal Papilla","volume":"142","author":"Shim","year":"2022","journal-title":"J. Investig. Dermatol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"118","DOI":"10.1007\/s11274-025-04336-4","article-title":"Keratinous bioresources: Their generation, microbial degradation, and value enhancement for biotechnological applications","volume":"41","author":"Vikash","year":"2025","journal-title":"World J. Microbiol. Biotechnol."},{"key":"ref_66","doi-asserted-by":"crossref","unstructured":"Lin, P.-Y., Stern, A., Peng, H.-H., Chen, J.-H., and Yang, H.-C. (2022). Redox and Metabolic Regulation of Intestinal Barrier Function and Associated Disorders. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms232214463"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"681","DOI":"10.1038\/s12276-023-00960-y","article-title":"Intestinal mucus components and secretion mechanisms: What we do and do not know","volume":"55","author":"Song","year":"2023","journal-title":"Exp. Mol. Med."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"1916376","DOI":"10.1080\/19490976.2021.1916376","article-title":"Healthy Intestinal Function Relies on Coordinated Enteric Nervous System, Immune System, and Epithelium Responses","volume":"13","author":"Kim","year":"2021","journal-title":"Gut Microbes"},{"key":"ref_69","doi-asserted-by":"crossref","unstructured":"Helenius, T.O., Antman, C.A., Asghar, M.N., Nystr\u00f6m, J.H., and Toivola, D.M. (2016). Keratins Are Altered in Intestinal Disease-Related Stress Responses. Cells, 5.","DOI":"10.3390\/cells5030035"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"2298","DOI":"10.1091\/mbc.E14-02-0736","article-title":"Keratin 8 absence down-regulates colonocyte HMGCS2 and modulates colonic ketogenesis and energy metabolism","volume":"26","author":"Helenius","year":"2015","journal-title":"Mol. Biol. Cell"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"1621","DOI":"10.1152\/physrev.00041.2019","article-title":"The Urothelium: Life in a Liquid Environment","volume":"100","author":"Dalghi","year":"2020","journal-title":"Physiol. Rev."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"1127","DOI":"10.1038\/s41385-022-00565-0","article-title":"The urothelium: A multi-faceted barrier against a harsh environment","volume":"15","author":"Jafari","year":"2022","journal-title":"Mucosal Immunol."},{"key":"ref_73","doi-asserted-by":"crossref","unstructured":"Peskar, D., Kos, M.K., Cerkvenik, U., Svete, A.N., and Erman, A. (2023). Sex-Dependent Differences in Blood-Urine Barrier Are Subtle but Significant in Healthy and Chronically Inflamed Mouse Bladders. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms242216296"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"631","DOI":"10.1016\/j.ajpath.2020.12.011","article-title":"Loss of Fibroblast Growth Factor Receptor 2 (FGFR2) Leads to Defective Bladder Urothelial Regeneration after Cyclophosphamide Injury","volume":"191","author":"Narla","year":"2021","journal-title":"Am. J. Pathol."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"982","DOI":"10.1038\/ncb3038","article-title":"Bladder cancers arise from distinct urothelial sub-populations","volume":"16","author":"Yamany","year":"2014","journal-title":"Nat. Cell Biol."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"1621","DOI":"10.1091\/mbc.E15-04-0230","article-title":"Sequential and compartmentalized action of Rabs, SNAREs, and MAL in the apical delivery of fusiform vesicles in urothelial umbrella cells","volume":"27","author":"Wankel","year":"2016","journal-title":"Mol. Biol. Cell"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"F1078","DOI":"10.1152\/ajprenal.00378.2023","article-title":"Keratin 5 basal cells are temporally regulated developmental and tissue repair progenitors in bladder urothelium","volume":"326","author":"Becknell","year":"2024","journal-title":"Am. J. Physiol. Renal. Physiol."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"469","DOI":"10.1016\/j.devcel.2013.07.017","article-title":"Retinoid signaling in progenitors controls specification and regeneration of the urothelium","volume":"26","author":"Gandhi","year":"2013","journal-title":"Dev. Cell"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"1336","DOI":"10.1038\/s41388-025-03311-5","article-title":"PTEN loss drives p53 LOH and immune evasion in a novel urothelial organoid model harboring p53 missense mutations","volume":"44","author":"Hamada","year":"2025","journal-title":"Oncogene"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"104798","DOI":"10.1016\/j.jmbbm.2021.104798","article-title":"Molecular origin of the effects of mutation on the structure and mechanical properties of human epithelial keratin K5\/K14","volume":"124","author":"Pan","year":"2021","journal-title":"J. Mech. Behav. Biomed. Mater."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"350","DOI":"10.1186\/s13054-024-05127-3","article-title":"Airway epithelium damage in acute respiratory distress syndrome","volume":"28","author":"Gerard","year":"2024","journal-title":"Crit. Care"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1111\/cea.13539","article-title":"Key role of the epithelium in chronic upper airways diseases","volume":"50","author":"Gohy","year":"2020","journal-title":"Clin. Exp. Allergy"},{"key":"ref_83","doi-asserted-by":"crossref","unstructured":"Raby, K.L., Michaeloudes, C., Tonkin, J., Chung, K.F., and Bhavsar, P.K. (2023). Mechanisms of airway epithelial injury and abnormal repair in asthma and COPD. Front. Immunol., 14.","DOI":"10.3389\/fimmu.2023.1201658"},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"240008","DOI":"10.1183\/16000617.0008-2024","article-title":"Neuroimmune recognition and regulation in the respiratory system","volume":"33","author":"Chen","year":"2024","journal-title":"Eur. Respir. Rev."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1172\/JCI150951","article-title":"Bitter taste signaling in tracheal epithelial brush cells elicits innate immune responses to bacterial infection","volume":"132","author":"Hollenhorst","year":"2022","journal-title":"J. Clin. Investig."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"213","DOI":"10.1186\/s12931-023-02514-2","article-title":"A comparative study of in vitro air-liquid interface culture models of the human airway epithelium evaluating cellular heterogeneity and gene expression at single cell resolution","volume":"24","author":"Prescott","year":"2023","journal-title":"Respir. Res."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1038\/onc.2010.456","article-title":"Keratins in health and cancer: More than mere epithelial cell markers","volume":"30","author":"Karantza","year":"2011","journal-title":"Oncogene"},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"521","DOI":"10.1042\/EBC20190017","article-title":"Keratin intermediate filaments: Intermediaries of epithelial cell migration","volume":"63","author":"Yoon","year":"2019","journal-title":"Essays Biochem."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"19385","DOI":"10.3390\/ijms141019385","article-title":"Keratin k15 as a biomarker of epidermal stem cells","volume":"14","author":"Bose","year":"2013","journal-title":"Int. J. Mol. Sci."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"252","DOI":"10.1111\/j.1440-1827.2011.02659.x","article-title":"Mixed squamous cell and glandular papilloma of the lung: A case study and literature review","volume":"61","author":"Inamura","year":"2011","journal-title":"Pathol. Int."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"777","DOI":"10.1002\/ijc.20942","article-title":"Real-time RT-PCR detection of CK19, CK7 and MUC1 mRNA for diagnosis of lymph node micrometastases in non small cell lung carcinoma","volume":"115","author":"Saintigny","year":"2005","journal-title":"Int. J. Cancer"},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"4397","DOI":"10.1007\/s00018-019-03424-7","article-title":"An intact keratin network is crucial for mechanical integrity and barrier function in keratinocyte cell sheets","volume":"77","author":"Karsch","year":"2020","journal-title":"Cell. Mol. Life Sci."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1016\/j.ceb.2014.12.008","article-title":"Keratins in health and disease","volume":"32","author":"Toivola","year":"2015","journal-title":"Curr. Opin. Cell Biol."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"201","DOI":"10.1038\/s41467-023-35795-8","article-title":"Control of protein stability by post-translational modifications","volume":"14","author":"Lee","year":"2023","journal-title":"Nat. Commun."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"102352","DOI":"10.1016\/j.sbi.2022.102352","article-title":"Uncovering post-translational modification-associated protein-protein interactions","volume":"74","author":"Wang","year":"2022","journal-title":"Curr. Opin. Struct. Biol."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"32","DOI":"10.1038\/s41392-022-01300-8","article-title":"Epigenetic and post-translational modifications in autophagy: Biological functions and therapeutic targets","volume":"8","author":"Shu","year":"2023","journal-title":"Signal Transduct. Target. Ther."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"329","DOI":"10.1038\/s41580-021-00441-y","article-title":"Modulation of cellular processes by histone and non-histone protein acetylation","volume":"23","author":"Shvedunova","year":"2022","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"bbad090","DOI":"10.1093\/bib\/bbad090","article-title":"Protein phosphorylation database and prediction tools","volume":"24","author":"Zhao","year":"2023","journal-title":"Brief. Bioinform."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"34","DOI":"10.1007\/s00018-023-05061-7","article-title":"Contributing roles of mitochondrial dysfunction and hepatocyte apoptosis in liver diseases through oxidative stress, post-translational modifications, inflammation, and intestinal barrier dysfunction","volume":"81","author":"LeFort","year":"2024","journal-title":"Cell. Mol. Life Sci."},{"key":"ref_100","doi-asserted-by":"crossref","unstructured":"Li, Y., Zhang, R., and Hei, H. (2023). Advances in post-translational modifications of proteins and cancer immunotherapy. Front. Immunol., 14.","DOI":"10.3389\/fimmu.2023.1229397"},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"639","DOI":"10.1021\/jacs.0c11566","article-title":"Controlling Kinase Activities by Selective Inhibition of Peptide Substrates","volume":"143","author":"Li","year":"2021","journal-title":"J. Am. Chem. Soc."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"5691","DOI":"10.1039\/D1CS00991E","article-title":"Dissecting the role of protein phosphorylation: A chemical biology toolbox","volume":"51","author":"Bilbrough","year":"2022","journal-title":"Chem. Soc. Rev."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"2190","DOI":"10.1016\/j.molcel.2022.05.007","article-title":"A journey from phosphotyrosine to phosphohistidine and beyond","volume":"82","author":"Hunter","year":"2022","journal-title":"Mol. Cell"},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"114907","DOI":"10.1016\/j.bcp.2022.114907","article-title":"Long non-coding RNAs in gastrointestinal cancers: Implications for protein phosphorylation","volume":"197","author":"Su","year":"2022","journal-title":"Biochem. Pharmacol."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"48","DOI":"10.1186\/s12943-018-0804-2","article-title":"Kinase-targeted cancer therapies: Progress, challenges and future directions","volume":"17","author":"Bhullar","year":"2018","journal-title":"Mol. Cancer"},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"986","DOI":"10.1016\/j.tibs.2024.08.004","article-title":"The expanding landscape of canonical and non-canonical protein phosphorylation","volume":"49","author":"Houles","year":"2024","journal-title":"Trends Biochem. Sci."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"102890","DOI":"10.1016\/j.sbi.2024.102890","article-title":"Exploring the conformational landscape of protein kinases","volume":"88","author":"Gough","year":"2024","journal-title":"Curr. Opin. Struct. Biol."},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"818","DOI":"10.1016\/j.tips.2019.09.002","article-title":"The Landscape of Atypical and Eukaryotic Protein Kinases","volume":"40","author":"Kanev","year":"2019","journal-title":"Trends Pharmacol. Sci."},{"key":"ref_109","doi-asserted-by":"crossref","unstructured":"Cipak, L. (2022). Protein Kinases: Function, Substrates, and Implication in Diseases. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms23073560"},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"177","DOI":"10.1038\/s41392-022-01038-3","article-title":"Targeting protein phosphatases for the treatment of inflammation-related diseases: From signaling to therapy","volume":"7","author":"Pan","year":"2022","journal-title":"Signal Transduct. Target. Ther."},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"4998","DOI":"10.1021\/acs.jctc.4c00206","article-title":"Effects of Phosphorylation on Protein Backbone Dynamics and Conformational Preferences","volume":"20","author":"Bickel","year":"2024","journal-title":"J. Chem. Theory Comput."},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"271","DOI":"10.3892\/ijmm.2017.3036","article-title":"The crucial role of protein phosphorylation in cell signaling and its use as targeted therapy (Review)","volume":"40","author":"Ardito","year":"2017","journal-title":"Int. J. Mol. Med."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"1814","DOI":"10.1016\/j.cell.2022.04.013","article-title":"mTOR substrate phosphorylation in growth control","volume":"185","author":"Battaglioni","year":"2022","journal-title":"Cell"},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"25316","DOI":"10.1021\/jacs.5c04275","article-title":"Phosphorylation-Inducing Molecules for Regulating Dynamic Cellular Processes","volume":"147","author":"Pergu","year":"2025","journal-title":"J. Am. Chem. Soc."},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1038\/s41575-020-00372-7","article-title":"Molecular and cellular mechanisms of liver fibrosis and its regression","volume":"18","author":"Kisseleva","year":"2021","journal-title":"Nat. Rev. Gastroenterol. Hepatol."},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1016\/j.mam.2018.09.002","article-title":"Liver fibrosis: Pathophysiology, pathogenetic targets and clinical issues","volume":"65","author":"Parola","year":"2019","journal-title":"Mol. Asp. Med."},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"633","DOI":"10.1038\/s41575-023-00807-x","article-title":"Hepatic inflammatory responses in liver fibrosis","volume":"20","author":"Hammerich","year":"2023","journal-title":"Nat. Rev. Gastroenterol. Hepatol."},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"397","DOI":"10.1038\/nrgastro.2017.38","article-title":"Mechanisms of hepatic stellate cell activation","volume":"14","author":"Tsuchida","year":"2017","journal-title":"Nat. Rev. Gastroenterol. Hepatol."},{"key":"ref_119","doi-asserted-by":"crossref","first-page":"583","DOI":"10.1038\/s41423-023-00983-5","article-title":"Mesenchymal stromal cells in hepatic fibrosis\/cirrhosis: From pathogenesis to treatment","volume":"20","author":"Yang","year":"2023","journal-title":"Cell. Mol. Immunol."},{"key":"ref_120","doi-asserted-by":"crossref","first-page":"912","DOI":"10.1053\/j.gastro.2020.11.051","article-title":"AGA Clinical Practice Update on Lifestyle Modification Using Diet and Exercise to Achieve Weight Loss in the Management of Nonalcoholic Fatty Liver Disease","volume":"160","author":"Younossi","year":"2021","journal-title":"Gastroenterology"},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"1639","DOI":"10.1002\/hep.21976","article-title":"Keratins let liver live: Mutations predispose to liver disease and crosslinking generates Mallory-Denk bodies","volume":"46","author":"Ku","year":"2007","journal-title":"Hepatology"},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"966","DOI":"10.1002\/hep.28493","article-title":"Keratins: Biomarkers and modulators of apoptotic and necrotic cell death in the liver","volume":"64","author":"Ku","year":"2016","journal-title":"Hepatology"},{"key":"ref_123","doi-asserted-by":"crossref","unstructured":"de Alteriis, G., Pugliese, G., Di Sarno, A., Muscogiuri, G., Barrea, L., Cossiga, V., Perruolo, G., Di Tolla, M.F., Zumbolo, F., and Formisano, P. (2023). Visceral Obesity and Cytokeratin-18 Antigens as Early Biomarkers of Liver Damage. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms241310885"},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"77","DOI":"10.14309\/ajg.0000000000001042","article-title":"Keratin-18: Diagnostic, Prognostic, and Theragnostic for Alcohol-Associated Hepatitis","volume":"116","author":"McClain","year":"2021","journal-title":"Am. J. Gastroenterol."},{"key":"ref_125","doi-asserted-by":"crossref","first-page":"3421","DOI":"10.1002\/hep4.2075","article-title":"Serum keratin-18 detects hepatic inflammation and predicts progression in compensated alcohol-associated liver disease","volume":"6","author":"Thorhauge","year":"2022","journal-title":"Hepatol. Commun."},{"key":"ref_126","doi-asserted-by":"crossref","first-page":"2046","DOI":"10.1016\/j.cgh.2019.11.050","article-title":"Keratin 18 Is a Diagnostic and Prognostic Factor for Acute Alcoholic Hepatitis","volume":"18","author":"Vatsalya","year":"2020","journal-title":"Clin. Gastroenterol. Hepatol."},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"144","DOI":"10.1002\/hep.32574","article-title":"PP2 protects from keratin mutation-associated liver injury and filament disruption via SRC kinase inhibition in male but not female mice","volume":"77","author":"Li","year":"2023","journal-title":"Hepatology"},{"key":"ref_128","doi-asserted-by":"crossref","first-page":"6063","DOI":"10.1073\/pnas.0936165100","article-title":"Keratin 8 and 18 mutations are risk factors for developing liver disease of multiple etiologies","volume":"100","author":"Ku","year":"2003","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"1858","DOI":"10.1002\/hep.27965","article-title":"PKC412 normalizes mutation-related keratin filament disruption and hepatic injury in mice by promoting keratin-myosin binding","volume":"62","author":"Kwan","year":"2015","journal-title":"Hepatology"},{"key":"ref_130","doi-asserted-by":"crossref","first-page":"1203","DOI":"10.1111\/liv.12608","article-title":"Keratins 8 and 18 are type II acute-phase responsive genes overexpressed in human liver disease","volume":"35","author":"Guldiken","year":"2015","journal-title":"Liver Int."},{"key":"ref_131","doi-asserted-by":"crossref","first-page":"161","DOI":"10.1186\/s13045-022-01381-6","article-title":"JMJD4-demethylated RIG-I prevents hepatic steatosis and carcinogenesis","volume":"15","author":"Li","year":"2022","journal-title":"J. Hematol. Oncol."},{"key":"ref_132","doi-asserted-by":"crossref","first-page":"eabc8433","DOI":"10.1126\/science.abc8433","article-title":"Structure-based decoupling of the pro- and anti-inflammatory functions of interleukin-10","volume":"371","author":"Saxton","year":"2021","journal-title":"Science"},{"key":"ref_133","doi-asserted-by":"crossref","first-page":"628","DOI":"10.1038\/s41586-024-07098-5","article-title":"IL-10 constrains sphingolipid metabolism to limit inflammation","volume":"627","author":"York","year":"2024","journal-title":"Nature"},{"key":"ref_134","doi-asserted-by":"crossref","first-page":"459","DOI":"10.1002\/hep.20277","article-title":"Keratin 8 and 18 hyperphosphorylation is a marker of progression of human liver disease","volume":"40","author":"Toivola","year":"2004","journal-title":"Hepatology"},{"key":"ref_135","doi-asserted-by":"crossref","first-page":"10751","DOI":"10.1096\/fj.201902476R","article-title":"PGC1\u03b1 protects against hepatic steatosis and insulin resistance via enhancing IL10-mediated anti-inflammatory response","volume":"34","author":"Wan","year":"2020","journal-title":"FASEB J."},{"key":"ref_136","doi-asserted-by":"crossref","first-page":"541","DOI":"10.1038\/s41568-021-00383-9","article-title":"The immunological and metabolic landscape in primary and metastatic liver cancer","volume":"21","author":"Li","year":"2021","journal-title":"Nat. Rev. Cancer"},{"key":"ref_137","doi-asserted-by":"crossref","first-page":"6","DOI":"10.1038\/s41572-020-00240-3","article-title":"Hepatocellular carcinoma","volume":"7","author":"Llovet","year":"2021","journal-title":"Nat. Rev. Dis. Primers"},{"key":"ref_138","doi-asserted-by":"crossref","first-page":"410","DOI":"10.1001\/jamasurg.2022.7989","article-title":"Management of Hepatocellular Carcinoma: A Review","volume":"158","author":"Brown","year":"2023","journal-title":"JAMA Surg."},{"key":"ref_139","doi-asserted-by":"crossref","first-page":"S228","DOI":"10.3350\/cmh.2024.0824","article-title":"Hepatocellular carcinoma: Updates on epidemiology, surveillance, diagnosis and treatment","volume":"31","author":"Hwang","year":"2025","journal-title":"Clin. Mol. Hepatol."},{"key":"ref_140","doi-asserted-by":"crossref","first-page":"1105","DOI":"10.1038\/s41417-024-00764-w","article-title":"Heterogeneity of hepatocellular carcinoma: From mechanisms to clinical implications","volume":"31","author":"Safri","year":"2024","journal-title":"Cancer Gene Ther."},{"key":"ref_141","doi-asserted-by":"crossref","first-page":"173","DOI":"10.1038\/s41575-024-01021-z","article-title":"Current and emerging strategies for the prevention of hepatocellular carcinoma","volume":"22","author":"Yeo","year":"2025","journal-title":"Nat. Rev. Gastroenterol. Hepatol."},{"key":"ref_142","doi-asserted-by":"crossref","first-page":"166335","DOI":"10.1016\/j.bbadis.2021.166335","article-title":"Deletion of kif3a in CK19 positive cells leads to primary cilia loss, biliary cell proliferation and cystic liver lesions in TAA-treated mice","volume":"1868","author":"Chen","year":"2022","journal-title":"Biochim. Biophys. Acta (BBA)\u2014Mol. Basis Dis."},{"key":"ref_143","doi-asserted-by":"crossref","first-page":"386","DOI":"10.5604\/16652681.1198813","article-title":"Cytokeratin19 positive hepatocellular carcinoma is associated with increased peritumoral ductular reaction","volume":"15","author":"Cai","year":"2016","journal-title":"Ann. Hepatol."},{"key":"ref_144","doi-asserted-by":"crossref","first-page":"540","DOI":"10.1002\/hep.24693","article-title":"Successful isolation of liver progenitor cells by aldehyde dehydrogenase activity in na\u00efve mice","volume":"55","author":"Best","year":"2012","journal-title":"Hepatology"},{"key":"ref_145","doi-asserted-by":"crossref","first-page":"674","DOI":"10.1136\/gutjnl-2012-304351","article-title":"Keratin 19: A key role player in the invasion of human hepatocellular carcinomas","volume":"63","author":"Govaere","year":"2014","journal-title":"Gut"},{"key":"ref_146","doi-asserted-by":"crossref","first-page":"64","DOI":"10.1186\/1476-4598-8-64","article-title":"JNK1 activation predicts the prognostic outcome of the human hepatocellular carcinoma","volume":"8","author":"Chang","year":"2009","journal-title":"Mol. Cancer"},{"key":"ref_147","doi-asserted-by":"crossref","first-page":"768","DOI":"10.7150\/ijbs.52279","article-title":"Linc-KILH potentiates Notch1 signaling through inhibiting K19 phosphorylation and promotes the malignancy of hepatocellular carcinoma","volume":"17","author":"Zhang","year":"2021","journal-title":"Int. J. Biol. Sci."},{"key":"ref_148","doi-asserted-by":"crossref","first-page":"e0204","DOI":"10.1097\/HC9.0000000000000204","article-title":"Epigenetic deregulation of MLF1 drives intrahepatic cholangiocarcinoma progression through EGFR\/AKT and Wnt\/\u03b2-catenin signaling","volume":"7","author":"Tang","year":"2023","journal-title":"Hepatol. Commun."},{"key":"ref_149","doi-asserted-by":"crossref","unstructured":"Van Haele, M., Moya, I.M., Karaman, R., Rens, G., Snoeck, J., Govaere, O., Nevens, F., Verslype, C., Topal, B., and Monbaliu, D. (2019). YAP and TAZ Heterogeneity in Primary Liver Cancer: An Analysis of Its Prognostic and Diagnostic Role. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20030638"},{"key":"ref_150","doi-asserted-by":"crossref","first-page":"239","DOI":"10.1159\/000490806","article-title":"Heterogeneity of Epigenetic and Epithelial Mesenchymal Transition Marks in Hepatocellular Carcinoma with Keratin 19 Proficiency","volume":"8","author":"Yokomichi","year":"2019","journal-title":"Liver Cancer"},{"key":"ref_151","doi-asserted-by":"crossref","first-page":"115519","DOI":"10.1016\/j.biopha.2023.115519","article-title":"Acetylation in pathogenesis: Revealing emerging mechanisms and therapeutic prospects","volume":"167","author":"Jiang","year":"2023","journal-title":"Biomed. Pharmacother."},{"key":"ref_152","doi-asserted-by":"crossref","first-page":"188837","DOI":"10.1016\/j.bbcan.2022.188837","article-title":"Acetyl-CoA regulates lipid metabolism and histone acetylation modification in cancer","volume":"1878","author":"He","year":"2023","journal-title":"Biochim. Biophys. Acta (BBA)\u2014Rev. Cancer"},{"key":"ref_153","doi-asserted-by":"crossref","first-page":"e326","DOI":"10.1038\/emm.2017.19","article-title":"Modulation of gene expression dynamics by co-transcriptional histone methylations","volume":"49","author":"Woo","year":"2017","journal-title":"Exp. Mol. Med."},{"key":"ref_154","doi-asserted-by":"crossref","first-page":"1245","DOI":"10.1016\/j.tplants.2023.06.006","article-title":"Multifaceted functions of histone deacetylases in stress response","volume":"28","author":"Cui","year":"2023","journal-title":"Trends Plant Sci."},{"key":"ref_155","doi-asserted-by":"crossref","first-page":"209","DOI":"10.1016\/j.semcancer.2021.03.001","article-title":"Targeting the acetylation signaling pathway in cancer therapy","volume":"85","author":"Dang","year":"2022","journal-title":"Semin. Cancer Biol."},{"key":"ref_156","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1186\/s13578-022-00821-7","article-title":"The phosphorylation to acetylation\/methylation cascade in transcriptional regulation: How kinases regulate transcriptional activities of DNA\/histone-modifying enzymes","volume":"12","author":"Zhao","year":"2022","journal-title":"Cell Biosci."},{"key":"ref_157","doi-asserted-by":"crossref","first-page":"90","DOI":"10.1038\/s41392-020-0196-9","article-title":"Regulating tumor suppressor genes: Post-translational modifications","volume":"5","author":"Chen","year":"2020","journal-title":"Signal Transduct. Target. Ther."},{"key":"ref_158","doi-asserted-by":"crossref","first-page":"e202006151","DOI":"10.1083\/jcb.202006151","article-title":"Lysine acetylation of cytoskeletal proteins: Emergence of an actin code","volume":"219","author":"A","year":"2020","journal-title":"J. Cell Biol."},{"key":"ref_159","doi-asserted-by":"crossref","first-page":"266","DOI":"10.1038\/s41392-024-01953-7","article-title":"Signaling pathways involved in colorectal cancer: Pathogenesis and targeted therapy","volume":"9","author":"Li","year":"2024","journal-title":"Signal Transduct. Target. Ther."},{"key":"ref_160","doi-asserted-by":"crossref","first-page":"589","DOI":"10.1016\/j.cell.2024.12.003","article-title":"Tumor \u201cage\u201d in early-onset colorectal cancer","volume":"188","author":"Mauri","year":"2025","journal-title":"Cell"},{"key":"ref_161","doi-asserted-by":"crossref","first-page":"609","DOI":"10.1038\/s41575-024-00934-z","article-title":"Mechanisms of metastatic colorectal cancer","volume":"21","author":"Sancho","year":"2024","journal-title":"Nat. Rev. Gastroenterol. Hepatol."},{"key":"ref_162","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1016\/j.semcdb.2021.08.010","article-title":"Colorectal keratins: Integrating nutrition, metabolism and colorectal health","volume":"128","author":"Evans","year":"2022","journal-title":"Semin. Cell Dev. Biol."},{"key":"ref_163","doi-asserted-by":"crossref","first-page":"876","DOI":"10.1038\/ncb2091","article-title":"Cytoskeletal keratin glycosylation protects epithelial tissue from injury","volume":"12","author":"Ku","year":"2010","journal-title":"Nat. Cell Biol."},{"key":"ref_164","doi-asserted-by":"crossref","first-page":"984","DOI":"10.1038\/cdd.2017.28","article-title":"Keratins regulate colonic epithelial cell differentiation through the Notch1 signalling pathway","volume":"24","author":"Misiorek","year":"2017","journal-title":"Cell Death Differ."},{"key":"ref_165","doi-asserted-by":"crossref","first-page":"3312","DOI":"10.1093\/nar\/gkf462","article-title":"Transcriptional deregulation of the keratin 18 gene in human colon carcinoma cells results from an altered acetylation mechanism","volume":"30","author":"Prochasson","year":"2002","journal-title":"Nucleic Acids Res."},{"key":"ref_166","doi-asserted-by":"crossref","first-page":"2964","DOI":"10.1101\/gad.8.24.2964","article-title":"Colorectal hyperplasia and inflammation in keratin 8-deficient FVB\/N mice","volume":"8","author":"Baribault","year":"1994","journal-title":"Genes Dev."},{"key":"ref_167","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1007\/s00018-021-04081-5","article-title":"Targeted deletion of keratin 8 in intestinal epithelial cells disrupts tissue integrity and predisposes to tumorigenesis in the colon","volume":"79","author":"Stenvall","year":"2021","journal-title":"Cell. Mol. Life Sci."},{"key":"ref_168","doi-asserted-by":"crossref","first-page":"456","DOI":"10.1186\/s12964-025-02422-6","article-title":"FUT11-Driven fucosylation coordinates K63 ubiquitination of keratin 17 to sustain psoriatic keratinocytes hyperproliferation","volume":"23","author":"Li","year":"2025","journal-title":"Cell Commun. Signal."}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/2\/972\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,1,20]],"date-time":"2026-01-20T11:30:50Z","timestamp":1768908650000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/2\/972"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,1,19]]},"references-count":168,"journal-issue":{"issue":"2","published-online":{"date-parts":[[2026,1]]}},"alternative-id":["ijms27020972"],"URL":"https:\/\/doi.org\/10.3390\/ijms27020972","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,1,19]]}}}