［1］NOMBELA P,MIGUEL-LOPEZ B,BLANCO S.The role of m6A,m5C and Ψ RNA modifications incancer:Novel therapeutic opportunities［J］.Mol Cancer,2021,20(1):18.
［2］EDELHEIT S,SCHWARTZ S,MUMBACH MR.Transcriptome-wide map ping of 5-methylcytidine RNA modifications in bacteria,archaea,and yeast reveals m5C within archaeal mRNAs［J］.PLoS Genet,2013,9(6)：e1003602.
［3］SQUIRES JE,PATEL HR,NOUSCH M,et al.Widespread occurrence of 5-methylcytosine in human coding and non-coding RNA［J］.Nucleic Acids Res,2012,40(11):5023-5033.
［4］BOHNSACK KE,HOBARTNER C,BOHNSACK MT.Eukaryotic 5-methylcytosine (m5C) RNA methyltransferases:Mechanisms,cellular functions,and links to disease［J］.Genes(Basel),2019,10(2):102.
［5］HONG B,BROCKENBROUGH JS,WU P,et al.Nop2p is required for pre-rRNA processing and 60S ribosome subunit synthesis in yeast［J］.Mol Cell Biol,1997,17(1):378-388.
［6］LIAO H,GAUR A,MCCONIE H,et al.Human NOP2/NSUN1 regulates ribosome biogenesis through non-catalytic complex formation with box C/D snoRNPs［J］.Nucleic Acids Res,2022,50(18):10695-10716.
［7］DELAUNAY S,PASCUAL G,FENG B,et al.Mitochondrial RNA modifications shape metabolic plasticity in metastasis［J］.Nature,2022,607(7919):593-603.
［8］METODIEV MD,SPAHR H,LOGUERCIO POLOSA P,et al.NSUN4 is a dual function mitochondrial protein required for both methylation of 12S rRNA and coordination of mitoribosomal assembly［J］.PLoS Genet,2014,10(2):e1004110.
［9］HEISSENBERGER C,LIENDL L,NAGELREITER F,et al.Loss of the ribosomal RNA methyltransferase NSUN5 impairs global protein synthesis and normal growth［J］.Nucleic Acids Res,2019,47(22):11807-11825.
［10］SELMI T,HUSSAIN S,DIETMANN S,et al.Sequence-and structure-specific cytosine-5 mRNA methylation by NSUN6［J］.Nucleic Acids Res,2021,49(2):1006-1022.
［11］LI J,LI H,LONG T,et al.Archaeal NSUN6 catalyzes m5C 72 modification on a wide-range of specific tRNAs［J］.Nucleic Acids Res,2019,47(4):2041-2055.
［12］AGUILO F,LI S,BALASUBRAMANIYAN N,et al.Deposition of 5-methylcytosine on enhancer RNAs enables the coactivator function of PGC-1α［J］.Cell Rep,2016,14(3):479-492.
［13］LI H,ZHU D,WU J,et al.New substrates and determinants for tRNA recognition of RNA methyltransferase DNMT2/TRDMT1［J］.RNA Biol,2021,18(12):2531-2545.
［14］SHEN H,ONTIVEROS RJ,OWENS MC,et al.TET-mediated 5-methylcytosine oxidation in tRNA promotes translation［J］.J Biol Chem,2021,296:100087.
［15］YIN X,HU L,XU Y.Structure and function of TET enzymes［J］.Adv Exp Med Biol,2022,1389:239-267.
［16］YANG X,YANG Y,SUN BF,et al.5-methylcytosine promotes mRNA export-NSUN2 as the methyltransferase and ALYREF as an m5C reader［J］.Cell Res,2017,27(5)：606-625.
［17］SU J,WU G,YE Y,et al.NSUN2-mediated RNA 5-methylcytosine promotes esophageal squamous cell carcinoma progression via LIN28B-dependent GRB2 mRNA stabilization［J］.Oncogene,2021,40(39):5814-5828.
［18］CHEN X,LI A.5-methylcytosine promotes pathogenesis of bladder cancer through stabilizing mRNAs［J］.Nat Cell Biol,2019,21(8):978-990.
［19］SUN Z,XUE S,ZHANG M,et al.Aberrant NSUN2-mediated m5C modification of H19 lncRNA is associated with poor differentiation of hepatocellular carcinoma［J］.Oncogene,2020,39(45):6906-6919.
［20］BLANCO S,DIETMANN S,FLORES JV,et al.Aberrant methylation of tRNAs links cellular stress to neuro-developmental disorders［J］.EMBO J,2014,33(18):2020-2039.
［21］SUN HY,HAO YJ,PING XL,et al.Research progress in dynamic RNA methylation modification and its regulation mech anism［J］.Life Science,2016,28(5):540-550.
［22］TUORTO F,HERBST F,ALERASOOL N,et al.The tRNA methyltransferase Dnmt2 is required for accurate polypeptide synthesis during haematopoiesis［J］.EMBO J,2015,34:2350-2362.
［23］FLORES JV,CORDERO-ESPINOZA L,OEZTUERK-WINDER F,et al.Cytosine-5 RNA methylation regulates neural stem cell differentiation and motility［J］.Stem Cell Reports,2017,8(1):112-124.
［24］JANIN M,ORTIZ-BARAHONA V,DE MOURA MC,et al.Epigenetic loss of RNA-methyltransferase NSUN5 in glioma targets ribosomes to drive a stress adaptive translational program［J］.Acta Neuropathol,2019,138(6):1053-1074.
［25］ZHANG M,SONG J,YUAN W,et al.Roles of RNA methylation on tumor immunity and clinical implications［J］.Front Immunol,2021,12:641507.
［26］JOHN SP,SUN J,CARLSON RJ,et al.IFIT1 exerts opposing regulatory effects on the inflammatory and interferon gene programs in LPS-activated human macrophages［J］.Cell Rep,2018,25(1):95-106.
［27］MENG S,ZHOU H,FENG Z,et al.Epigenetics in neurodevelopment:emerging role of circular RNA［J］.Front Cell Neurosci,2019,13:327.
［28］STEPANOV G,ZHURAVLEV E,SHENDER V,et al.Nucleotide modifications decrease innate immune response induced by synthetic analogs of snRNAs and snoRNAs［J］.Genes(Basel),2018,9(11):531.
［29］HOERNES TP,CLEMENTI N,FASERL K,et al.Nucleotide modifications within bacterial messenger RNAs regulate their translation and are able to rewire the genetic code［J］.Nucleic Acids Research,2016,44(2):852-862.
［30］MEI L,SHEN C,MIAO R,et al.RNA methyltransferase NSUN2 promotes gastric cancer cell proliferation by repressing p57Kip2 by an m5C-dependent manner［J］.Cell Death Dis,2020,11(4):270.
［31］JIANG Z,LI S,HAN MJ,et al.High expression of NSUN5 promotes cell proliferation via cell cycle regulation in colorectal cancer［J］.Am J Transl Res,2020,12(7):3858-3870.
［32］YANG R,LIANG X,WANG H,et al.The RNA methyltransferase NSUN6 suppresses pancreatic cancer development by regulating cell proliferation［J］.EBio Medicine,2021,63:103195.
［33］WANG J,LI Y,XU B,et al.ALYREF drives cancer cell proliferation through an ALYREF-MYC positive feedback loop in glioblastoma［J］.Onco Targets Ther,2021,14:145-155.
［34］HE Y,YU X,LI J,et al.Role of m5C-related regulatory genes in the diagnosis and prognosis of hepatocellular carcinoma［J］.Am J Transl Res,2020,12(3):912-922.
［35］XUE C,ZHAO Y,LI G,et al.Multi-omic analyses of the m5C regulator ALYREF reveal its essential roles in hepatocellular carcinoma［J］.Front Oncol,2021,11:633415.
［36］LI Y,LI J,LUO M,et al.Novel long noncoding RNA NMR promotes tumor progression via NSUN2 and BPTF in esophageal squamous cell carcinoma［J］.Cancer Lett,2018,430:57-66.
［37］WANG L,ZHANG J,SU Y,et al.Distinct roles of m5C RNA methyltransferase NSUN2 in major gynecologic cancers［J］.Front Oncol,2022,12:786266.
［38］BI J,HUANG Y,LIU Y.Effect of NOP2 knockdown on colon cancer cell proliferation,migration,and invasion［J］.Transl Cancer Res,2019,8(6):2274-2283.
［39］SUN F,WU K,YAO Z,et al.Long noncoding RNA PVT1 promotes prostate cancer metastasis by increasing NOP2 expression via targeting tumor suppressor microRNAs［J］.Onco Targets Ther,2020,13:6755-6765.
［40］SUN F,WU K,YAO Z,et al.Long noncoding RNA LINC00963 induces NOP2 expression by sponging tumor suppressor miR-542-3p to promote metastasis in prostate cancer［J］.Aging (Albany NY),2020,12(12):11500-11516.
［41］AWAH CU,WINTER J,MAZDOOM CM,et al.NSUN6,an RNA methyltransferase of 5-mC controls glioblastoma response to temozolomide (TMZ) via NELFB and RPS6KB2 interaction［J］.Cancer Biol Ther,2021,22(10-12):587-597.
［42］LI W,XU L.Epigenetic function of TET family,5-methylcytosine,and 5-hydroxymethylcytosine in hematologic malignancies［J］.Oncol Res Treat,2019,42(6):309-318.
［43］JUNG HA,JUNG CW,JANG JH.Mutations in genes affecting DNA methylation enhances responses to decitabine in patients with myelodysplastic syndrome［J］.Korean J Intern Med,2021,36(2):413-423.
［44］REILLY BM,LUGER T,PARK S,et al.5-azacytidine transiently restores dysregulated erythroid differentiation gene expression in TET2-deficient erythroleukemia cells［J］.Mol Cancer Res,2021,19(3):451-464.
［45］XUE M,SHI Q,ZHENG L,et al.Gene signatures of m5C regulators may predict prognoses of patients with head and neck squamous cell carcinoma［J］.Am J Transl Res,2020,12(10):6841-6852.
［46］WANG JZ,ZHU W,HAN J,et al.The role of the HIF-1α/ALYREF/PKM2 axis in glycolysis and tumorigenesis of bladder cancer［J］.Cancer Commun (Lond),2021,41(7):560-575.
［47］GAO Y,WANG Z,ZHU Y,et al.NOP2/Sun RNA methyltransferase 2 promotes tumor progression via its interacting partner RPL6 in gallbladder carcinoma［J］.Cancer Sci,2019,110(11):3510-3519.
［48］PAN J,HUANG Z,XU Y.m5C RNA methylation regulators predict prognosis and regulate the immune microenvironment in lung squamous cell carcinoma［J］.Front Oncol,2021,11:657466.
［49］YANG JC,RISCH E,ZHANG M,et al.Association of tRNA methyltransferase NSUN2/IGF-Ⅱ molecular signature with ovarian cancer survival［J］.Future Oncol,2017,13(22):1981-1990.
［50］HUANG Z,PAN J,WANG H,et al.Prognostic significance and tumor immune microenvironment heterogenicity of m5C RNA methylation regulators in triple-negative breast cancer［J］.Front Cell Dev Biol,2021,9:657547.
［51］LU L,ZHU G,ZENG H,et al.High tRNA transferase NSUN2 gene expression is associated with poor prognosis in head and neck squamous carcinoma［J］.Cancer Invest,2018,36(4):246-253.
［52］KAR SP,BEESLEY J,AMIN AL OLAMA A,et al.Genome-wide Meta-analyses of breast,ovarian,and prostate cancer association studies identify multiple new susceptibility loci shared by at least two cancer types［J］.Cancer Discov,2016,6(9):1052-1067.
［53］LI H,JIANG H,HUANG Z,et al.Prognostic value of an m5C RNA methylation regulator-related signature for clear cell renal cell carcinoma［J］.Cancer Manag Res,2021,13:6673-6687.
［54］HUANG M,ZHANG Y,OU X,et al.m5C-related signatures for predicting prognosis in cutaneous melanoma with machine learning［J］.J Oncol,2021,2021:6173206.
［55］SATO K,TAHATA K,AKIMOTO K.Five genes associated with survival in patients with lower-grade gliomas were identified by information-theoretical analysis［J］.Anticancer Res,2020,40(5):2777-2785.
［56］ELHARDT W,SHANMUGAM R,JURKOWSKI TP,et al.Somatic cancer mutations in the DNMT2 tRNA methyltransferase alter its catalytic properties［J］.Biochimie,2015,112:66-72.
［57］TAKAI H,MASUDA K,SATO T,et al.5-Hydroxymethylcytosine plays a critical role in glioblastomagenesis by recruiting the CHTOP methylosome complex［J］.Cell Rep,2014,9:48-60.
［58］GARCIA MG,CARELLA A,URDINGUIO RG,et al.Epigenetic dysregulation of TET2 in human glioblastoma［J］.Oncotarget,2018,9(40):25922-25934.
［59］CARELLA A,TEJEDOR JR,GARCIA MG,et al.Epigenetic downregulation of TET3 reduces genome-wide 5-hmC levels and promotes glioblastoma tumorigenesis［J］.Int J Cancer,2020,146(2):373-387.
［60］SCHAEFER M,HAGEMANN S,HANNA K,et al.Azacytidine inhibits RNA methylation at DNMT2 target sites in human cancer cell lines［J］.Cancer Res,2009,69(20):8127-8132.