长链非编码RNA在调控抗病毒天然免疫应答中的研究进展
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摘要
天然免疫系统是机体防御病原微生物入侵的第一道防线,其生物效应依赖于复杂的信号传导和动态的基因表达调控。长链非编码RNA(long non-coding RNA,lncRNA)是一类不编码蛋白质的RNA转录本。近年来,越来越多的研究表明lncRNA具有在表观遗传、转录、转录后及翻译等水平调控基因表达的能力,其功能异常与多种疾病的发生发展密切相关。本文综述了病毒和宿主编码lncRNA参与调控抗病毒天然免疫应答的研究进展,广泛深入地开展相关研究不仅有助于理解病毒与宿主间的相互作用,而且有益于发现新的抗病毒策略。
The innate immune system is the first line of defense against microbial pathogens. The innate immune response relies on a highly integrated and dynamic gene-expression program controlled by transcriptional and post-transcriptional mechanisms. In recent years,accumulating data have suggested that long non-coding RNAs(lncRNAs),an important class of regulatory RNAs,have powerful abilities to regulate gene expression. These abilities range from epigenetics,transcription,RNA processing,post-transcriptional regulation,and protein translation to protein translocation. These actions are involved in a diverse range of biologic processes,including normal physiologic processes,cell development,viral infection,the immune response and human diseases.This review discussed recent findings on the emerging roles of lncRNA in controlling innate antiviral immunity and their mechanisms of action. Gaining better understanding of these processes could facilitate the development of novel therapeutics.
The innate immune system is the first line of defense against microbial pathogens. The innate immune response relies on a highly integrated and dynamic gene-expression program controlled by transcriptional and post-transcriptional mechanisms. In recent years,accumulating data have suggested that long non-coding RNAs(lncRNAs),an important class of regulatory RNAs,have powerful abilities to regulate gene expression. These abilities range from epigenetics,transcription,RNA processing,post-transcriptional regulation,and protein translation to protein translocation. These actions are involved in a diverse range of biologic processes,including normal physiologic processes,cell development,viral infection,the immune response and human diseases.This review discussed recent findings on the emerging roles of lncRNA in controlling innate antiviral immunity and their mechanisms of action. Gaining better understanding of these processes could facilitate the development of novel therapeutics.
引文
[1]Hangauer M J,Vaughn I W,McManus M T.Pervasive transcription of the human genome produces thousands of previously unidentified long intergenic noncoding RNAs[J/OL].PLoS Genet,2013,9(6):e1003569.
[2]Brown C J,Ballabio A,Rupert J L,Lafreniere R G,Grompe M,Tonlorenzi R,Willard H F.A gene from the region of the human X inactivation centre is expressed exclusively from the inactive X chromosome[J].Nature,1991,349(6304):38-44.
[3]Derrien T,Johnson R,Bussotti G,Tanzer A,Djebali S,Tilgner H,Guernec G,Martin D,Merkel A,Knowles D G,Lagarde J,Veeravalli L,Ruan X,Ruan Y,Lassmann T,Carninci P,Brown J B,Lipovich L,Gonzalez J M,Thomas M,Davis C A,Shiekhattar R,Gingeras T R,Hubbard T J,Notredame C,Harrow J,Guigo R.The GENCODE v7 catalog of human long noncoding RNAs:analysis of their gene structure,evolution,and expression[J].Genome Res,2012,22(9):1775-1789.
[4]Djebali S,Davis C A,Merkel A,Dobin A,Lassmann T,Mortazavi A,Tanzer A,Lagarde J,Lin W,Schlesinger F,Xue C,Marinov G K,Khatun J,Williams B A,Zaleski C,Rozowsky J,Roder M,Kokocinski F,Abdelhamid R F,Alioto T,Antoshechkin I,Baer M T,Bar N S,Batut P,Bell K,Bell I,Chakrabortty S,Chen X,Chrast J,Curado J,Derrien T,Drenkow J,Dumais E,Dumais J,Duttagupta R,Falconnet E,Fastuca M,Fejes-Toth K,Ferreira P,Foissac S,Fullwood M J,Gao H,Gonzalez D,Gordon A,Gunawardena H,Howald C,Jha S,Johnson R,Kapranov P,King B,Kingswood C,Luo O J,Park E,Persaud K,Preall J B,Ribeca P,Risk B,Robyr D,Sammeth M,Schaffer L,See L-H,Shahab A,Skancke J,Suzuki A M,Takahashi H,Tilgner H,Trout D,Walters N,Wang H,Wrobel J,Yu Y,Ruan X,Hayashizaki Y,Harrow J,Gerstein M,Hubbard T,Reymond A,Antonarakis S E,Hannon G,Giddings M C,Ruan Y,Wold B,Carninci P,Guigo R,Gingeras T R.Landscape of transcription in human cells[J].Nature,2012,489(7414):101-108.
[5]Zhao J,Sun B K,Erwin J A,Song J J,Lee J T.Polycomb proteins targeted by a short repeat RNA to the mouse X chromosome[J].Science,2008,322(5902):750-756.
[6]Martianov I,Ramadass A,Barros A S,Chow N,Akoulitchev A.Repression of the human dihydrofolate reductase gene by a non-coding interfering transcript[J].Nature,2007,445(7128):666-670.
[7]Cesana M,Cacchiarelli D,Legnini I,Santini T,Sthandier O,Chinappi M,Tramontano A,Bozzoni I.Along noncoding RNA controls muscle differentiation by functioning as a competing endogenous RNA[J].Cell,2011,147(2):358-369.
[8]Faghihi M A,Modarresi F,Khalil A M,Wood D E,Sahagan B G,Morgan T E,Finch C E,Laurent G S,Kenny P J,Wahlestedt C.Expression of a noncoding RNA is elevated in Alzheimer′s disease and drives rapid feed-forward regulation of beta-secretase[J].Nat Med,2008,14(7):723-730.
[9]Carrieri C,Cimatti L,Biagioli M,Beugnet A,Zucchelli S,Fedele S,Pesce E,Ferrer I,Collavin L,Santoro C,Forrest A R R,Carninci P,Biffo S,Stupka E,Gustincich S.Long non-coding antisense RNAcontrols Uchl1 translation through an embedded SINEB2 repeat[J].Nature,2012,491(7424):454-457.
[10]Yoon J H,Abdelmohsen K,Srikantan S,Yang X L,Martindale J L,De S,Huarte M,Zhan M,Becker KG,Gorospe M.LincRNA-p21 suppresses target mRNAtranslation[J].Mol Cell,2012,47(4):648-655.
[11]Hamonic G,Pasternak J A,Wilson H L.Recognizing conserved non-canonical localization patterns of toll-like receptors in tissues and across species[J].Cell Tissue Res,2018,372(1):1-11.
[12]Ma H,Han P,Ye W,Chen H,Zheng X,Cheng L,Zhang L,Yu L,Wu X a,Xu Z,Lei Y,Zhang F.The long noncoding RNA NEAT1 exerts antihantaviral effects by acting as positive feedback for RIG-I signaling[J/OL].J Virol,2017,91(9):e02250-02216.
[13]Miyashita M,Oshiumi H,Matsumoto M,Seya T.DDX60,a DEXD/H box helicase,is a novel antiviral factor promoting RIG-I-like receptor-mediated signaling[J].Mol Cell Biol,2011,31(18):3802-3819.
[14]Morchikh M,Cribier A,Raffel R,Amraoui S,Cau J,Severac D,Dubois E,Schwartz O,Bennasser Y,Benkirane M.HEXIM1 and NEAT1 long non-coding RNA form a multi-subunit complex that regulates DNA-mediated innate immune response[J].Mol Cell,2017,67(3):387-399.
[15]Penkala I,Wang J,Syrett C M,Goetzl L,Lopez C B,Anguera M C.lncRHOXF1,a long noncoding RNAfrom the X chromosome that suppresses viral response genes during development of the early human placenta[J].Mol Cell Biol,(2016),36(12):1764-1775.
[16]Jiang M,Zhang S,Yang Z,Lin H,Zhu J,Liu L,Wang W,Liu S,Liu W,Ma Y,Zhang L,Cao X.Selfrecognition of an inducible host lncRNA by RIG-Ifeedback restricts innate immune response[J].Cell,2018,173(4):906-919.
[17]Xie Q,Chen S,Tian R,Huang X,Deng R,Xue B,Qin Y,Xu Y,Wang J,Guo M,Chen J,Tang S,Li G,Zhu H.Long noncoding RNA ITPRIP-1 positively regulates the innate immune response through promotion of oligomerization and activation of MDA[J/OL].JVirol,2018,92(17):e00507-00518.
[18]Gomez J A,Wapinski O L,Yang Y W,Bureau J-F,Gopinath S,Monack D M,Chang H Y,Brahic M,Kirkegaard K.The NeST long ncRNA controls microbial susceptibility and epigenetic activation of the interferon-gamma locus[J].Cell,2013,152(4):743-754.
[19]Ouyang J,Zhu X,Chen Y,Wei H,Chen Q,Chi X,Qi B,Zhang L,Zhao Y,Gao G F,Wang G,Chen J-L.NRAV,a long noncoding RNA,modulates antiviral responses through suppression of interferon-stimulated gene transcription[J].Cell Host Microbe,2014,16(5):616-626.
[20]Nishitsuji H,Ujino S,Yoshio S,Sugiyama M,Mizokami M,Kanto T,Shimotohno K.Long noncoding RNA#32 contributes to antiviral responses by controlling interferon-stimulated gene expression[J].Proc Natl Acad Sci U S A,2016,113(37):10388-10393.
[21]Carpenter S,Aiello D,Atianand M K,Ricci E P,Gandhi P,Hall L L,Byron M,Monks B,Henry-Bezy M,Lawrence J B,O′Neill L A J,Moore M J,Caffrey D R,Fitzgerald K A.A long noncoding RNA mediates both activation and repression of immune response genes[J].Science,2013,341(6147):789-792.
[22]Atianand M K,Hu W,Satpathy A T,Shen Y,Ricci EP,Alvarez-Dominguez J R,Bhatta A,Schattgen S A,McGowan J D,Blin J,Braun J E,Gandhi P,Moore MJ,Chang H Y,Lodish H F,Caffrey D R,Fitzgerald KA.A long noncoding RNA lincRNA-EPS acts as a transcriptional brake to restrain inflammation[J].Cell,2016,165(7):1672-1685.
[23]Kambara H,Niazi F,Kostadinova L,Moonka D K,Siegel C T,Post A B,Carnero E,Barriocanal M,Fortes P,Anthony D D,Valadkhan S.Negative regulation of the interferon response by an interferoninduced long non-coding RNA[J].Nucleic Acids Res,2014,42(16):10668-10680.
[24]Carnero E,Barriocanal M,Prior C,Pablo Unfried J,Segura V,Guruceaga E,Enguita M,Smerdou C,Gastaminza P,Fortes P.Long noncoding RNA EGOTnegatively affects the antiviral response and favors HCVreplication[J].EMBO Rep,2016,17(7):1013-1028.
[25]Barriocanal M,Carnero E,Segura V,Fortes P.Long non-coding RNA BST2/BISPR is induced by IFN and regulates the expression of the antiviral factor tetherin[J].Front Immunol,2014,5:655.
[26]Rapicavoli N A,Qu K,Zhang J,Mikhail M,Laberge R-M,Chang H Y.A mammalian pseudogene lncRNAat the interface of inflammation and anti-inflammatory therapeutics[J/OL].Elife,2013,2:e00762.
[27]Liu B,Sun L,Liu Q,Gong C,Yao Y,Lv X,Lin L,Yao H,Su F,Li D,Zeng M,Song E.A cytoplasmic NF-kappaB interacting long noncoding RNA blocks IkappaB phosphorylation and suppresses breast cancer metastasis[J].Cancer Cell,2015,27(3):370-381.
[28]Krawczyk M,Emerson B M.p50-associated COX-2extragenic RNA(PACER)activates COX-2 gene expression by occluding repressive NF-kappaBcomplexes[J/OL].Elife,2014,3:e01776.
[29]Li Z,Chao T-C,Chang K-Y,Lin N,Patil V S,Shimizu C,Head S R,Burns J C,Rana T M.The long noncoding RNA THRIL regulates TNFalpha expression through its interaction with hnRNPL[J].Proc Natl Acad Sci U S A,2014,111(3):1002-1007.
[30]Iiott N E,Heward J A,Roux B,Tsitsiou E,Fenwick P S,Lenzi L,Goodhead I,Hertz-Fowler C,Heger A,Hall N,Donnelly L E,Sims D,Lindsay M A.Long non-coding RNAs and enhancer RNAs regulate the lipopolysaccharide-induced inflammatory response in human monocytes[J].Nat Commun,2014,5:3979.
[31]Lu J,Wu X,Hong M,Tobias P,Han J.A potential suppressive effect of natural antisense IL-1beta RNA on lipopolysaccharide-induced IL-1beta expression[J].JImmunol,2013,190(12):6570-6578.
[32]Mathews M B.Genes for VA-RNA in adenovirus 2[J].Cell,1975,6(2):223-229.
[33]Rossetto C C,Pari G S.Kaposi′s sarcoma-associated herpesvirus noncoding polyadenylated nuclear RNAinteracts with virus-and host cell-encoded proteins and suppresses expression of genes involved in immune modulation[J].J Virol,2011,85(24):13290-13297.
[34]Rossetto C C,Tarrant-Elorza M,Verma S,Purushothaman P,Pari G S.Regulation of viral and cellular gene expression by Kaposi′s sarcoma-associated herpesvirus polyadenylated nuclear RNA[J].J Virol,2013,87(10):5540-5553.
[35]Bidet K,Dadlani D,Garcia-Blanco M A.G3BP1,G3BP2 and CAPRIN1 are required for translation of interferon stimulated mRNAs and are targeted by a dengue virus non-coding RNA[J/OL].PLoS Pathog,2014,10(7):e1004242.
[36]Chang R-Y,Hsu T-W,Chen Y-L,Liu S-F,Tsai Y-J,Lin Y-T,Chen Y-S,Fan Y-H.Japanese encephalitis virus non-coding RNA inhibits activation of interferon by blocking nuclear translocation of interferon regulatory factor 3[J].Vet Microbiol,2013,166(1-2):11-21.
[37]Schuessler A,Funk A,Lazear H M,Cooper D A,Torres S,Daffis S,Jha B K,Kumagai Y,Takeuchi O,Hertzog P,Silverman R,Akira S,Barton D J,Diamond M S,Khromykh A A.West Nile virus noncoding subgenomic RNA contributes to viral evasion of the type I interferon-mediated antiviral response[J].JVirol,2012,86(10):5708-5718.
[38]Mathews M B.Structure,function,and evolution of adenovirus virus-associated RNAs[J].Curr Top Microbiol Immunol,1995,199(Pt 2):173-187.
[39]Nanbo A,Inoue K,Adachi-Takasawa K,Takada K.Epstein-Barr virus RNA confers resistance to interferonalpha-induced apoptosis in Burkitt′s lymphoma[J].Embo J,2002,21(5):954-965.
[2]Brown C J,Ballabio A,Rupert J L,Lafreniere R G,Grompe M,Tonlorenzi R,Willard H F.A gene from the region of the human X inactivation centre is expressed exclusively from the inactive X chromosome[J].Nature,1991,349(6304):38-44.
[3]Derrien T,Johnson R,Bussotti G,Tanzer A,Djebali S,Tilgner H,Guernec G,Martin D,Merkel A,Knowles D G,Lagarde J,Veeravalli L,Ruan X,Ruan Y,Lassmann T,Carninci P,Brown J B,Lipovich L,Gonzalez J M,Thomas M,Davis C A,Shiekhattar R,Gingeras T R,Hubbard T J,Notredame C,Harrow J,Guigo R.The GENCODE v7 catalog of human long noncoding RNAs:analysis of their gene structure,evolution,and expression[J].Genome Res,2012,22(9):1775-1789.
[4]Djebali S,Davis C A,Merkel A,Dobin A,Lassmann T,Mortazavi A,Tanzer A,Lagarde J,Lin W,Schlesinger F,Xue C,Marinov G K,Khatun J,Williams B A,Zaleski C,Rozowsky J,Roder M,Kokocinski F,Abdelhamid R F,Alioto T,Antoshechkin I,Baer M T,Bar N S,Batut P,Bell K,Bell I,Chakrabortty S,Chen X,Chrast J,Curado J,Derrien T,Drenkow J,Dumais E,Dumais J,Duttagupta R,Falconnet E,Fastuca M,Fejes-Toth K,Ferreira P,Foissac S,Fullwood M J,Gao H,Gonzalez D,Gordon A,Gunawardena H,Howald C,Jha S,Johnson R,Kapranov P,King B,Kingswood C,Luo O J,Park E,Persaud K,Preall J B,Ribeca P,Risk B,Robyr D,Sammeth M,Schaffer L,See L-H,Shahab A,Skancke J,Suzuki A M,Takahashi H,Tilgner H,Trout D,Walters N,Wang H,Wrobel J,Yu Y,Ruan X,Hayashizaki Y,Harrow J,Gerstein M,Hubbard T,Reymond A,Antonarakis S E,Hannon G,Giddings M C,Ruan Y,Wold B,Carninci P,Guigo R,Gingeras T R.Landscape of transcription in human cells[J].Nature,2012,489(7414):101-108.
[5]Zhao J,Sun B K,Erwin J A,Song J J,Lee J T.Polycomb proteins targeted by a short repeat RNA to the mouse X chromosome[J].Science,2008,322(5902):750-756.
[6]Martianov I,Ramadass A,Barros A S,Chow N,Akoulitchev A.Repression of the human dihydrofolate reductase gene by a non-coding interfering transcript[J].Nature,2007,445(7128):666-670.
[7]Cesana M,Cacchiarelli D,Legnini I,Santini T,Sthandier O,Chinappi M,Tramontano A,Bozzoni I.Along noncoding RNA controls muscle differentiation by functioning as a competing endogenous RNA[J].Cell,2011,147(2):358-369.
[8]Faghihi M A,Modarresi F,Khalil A M,Wood D E,Sahagan B G,Morgan T E,Finch C E,Laurent G S,Kenny P J,Wahlestedt C.Expression of a noncoding RNA is elevated in Alzheimer′s disease and drives rapid feed-forward regulation of beta-secretase[J].Nat Med,2008,14(7):723-730.
[9]Carrieri C,Cimatti L,Biagioli M,Beugnet A,Zucchelli S,Fedele S,Pesce E,Ferrer I,Collavin L,Santoro C,Forrest A R R,Carninci P,Biffo S,Stupka E,Gustincich S.Long non-coding antisense RNAcontrols Uchl1 translation through an embedded SINEB2 repeat[J].Nature,2012,491(7424):454-457.
[10]Yoon J H,Abdelmohsen K,Srikantan S,Yang X L,Martindale J L,De S,Huarte M,Zhan M,Becker KG,Gorospe M.LincRNA-p21 suppresses target mRNAtranslation[J].Mol Cell,2012,47(4):648-655.
[11]Hamonic G,Pasternak J A,Wilson H L.Recognizing conserved non-canonical localization patterns of toll-like receptors in tissues and across species[J].Cell Tissue Res,2018,372(1):1-11.
[12]Ma H,Han P,Ye W,Chen H,Zheng X,Cheng L,Zhang L,Yu L,Wu X a,Xu Z,Lei Y,Zhang F.The long noncoding RNA NEAT1 exerts antihantaviral effects by acting as positive feedback for RIG-I signaling[J/OL].J Virol,2017,91(9):e02250-02216.
[13]Miyashita M,Oshiumi H,Matsumoto M,Seya T.DDX60,a DEXD/H box helicase,is a novel antiviral factor promoting RIG-I-like receptor-mediated signaling[J].Mol Cell Biol,2011,31(18):3802-3819.
[14]Morchikh M,Cribier A,Raffel R,Amraoui S,Cau J,Severac D,Dubois E,Schwartz O,Bennasser Y,Benkirane M.HEXIM1 and NEAT1 long non-coding RNA form a multi-subunit complex that regulates DNA-mediated innate immune response[J].Mol Cell,2017,67(3):387-399.
[15]Penkala I,Wang J,Syrett C M,Goetzl L,Lopez C B,Anguera M C.lncRHOXF1,a long noncoding RNAfrom the X chromosome that suppresses viral response genes during development of the early human placenta[J].Mol Cell Biol,(2016),36(12):1764-1775.
[16]Jiang M,Zhang S,Yang Z,Lin H,Zhu J,Liu L,Wang W,Liu S,Liu W,Ma Y,Zhang L,Cao X.Selfrecognition of an inducible host lncRNA by RIG-Ifeedback restricts innate immune response[J].Cell,2018,173(4):906-919.
[17]Xie Q,Chen S,Tian R,Huang X,Deng R,Xue B,Qin Y,Xu Y,Wang J,Guo M,Chen J,Tang S,Li G,Zhu H.Long noncoding RNA ITPRIP-1 positively regulates the innate immune response through promotion of oligomerization and activation of MDA[J/OL].JVirol,2018,92(17):e00507-00518.
[18]Gomez J A,Wapinski O L,Yang Y W,Bureau J-F,Gopinath S,Monack D M,Chang H Y,Brahic M,Kirkegaard K.The NeST long ncRNA controls microbial susceptibility and epigenetic activation of the interferon-gamma locus[J].Cell,2013,152(4):743-754.
[19]Ouyang J,Zhu X,Chen Y,Wei H,Chen Q,Chi X,Qi B,Zhang L,Zhao Y,Gao G F,Wang G,Chen J-L.NRAV,a long noncoding RNA,modulates antiviral responses through suppression of interferon-stimulated gene transcription[J].Cell Host Microbe,2014,16(5):616-626.
[20]Nishitsuji H,Ujino S,Yoshio S,Sugiyama M,Mizokami M,Kanto T,Shimotohno K.Long noncoding RNA#32 contributes to antiviral responses by controlling interferon-stimulated gene expression[J].Proc Natl Acad Sci U S A,2016,113(37):10388-10393.
[21]Carpenter S,Aiello D,Atianand M K,Ricci E P,Gandhi P,Hall L L,Byron M,Monks B,Henry-Bezy M,Lawrence J B,O′Neill L A J,Moore M J,Caffrey D R,Fitzgerald K A.A long noncoding RNA mediates both activation and repression of immune response genes[J].Science,2013,341(6147):789-792.
[22]Atianand M K,Hu W,Satpathy A T,Shen Y,Ricci EP,Alvarez-Dominguez J R,Bhatta A,Schattgen S A,McGowan J D,Blin J,Braun J E,Gandhi P,Moore MJ,Chang H Y,Lodish H F,Caffrey D R,Fitzgerald KA.A long noncoding RNA lincRNA-EPS acts as a transcriptional brake to restrain inflammation[J].Cell,2016,165(7):1672-1685.
[23]Kambara H,Niazi F,Kostadinova L,Moonka D K,Siegel C T,Post A B,Carnero E,Barriocanal M,Fortes P,Anthony D D,Valadkhan S.Negative regulation of the interferon response by an interferoninduced long non-coding RNA[J].Nucleic Acids Res,2014,42(16):10668-10680.
[24]Carnero E,Barriocanal M,Prior C,Pablo Unfried J,Segura V,Guruceaga E,Enguita M,Smerdou C,Gastaminza P,Fortes P.Long noncoding RNA EGOTnegatively affects the antiviral response and favors HCVreplication[J].EMBO Rep,2016,17(7):1013-1028.
[25]Barriocanal M,Carnero E,Segura V,Fortes P.Long non-coding RNA BST2/BISPR is induced by IFN and regulates the expression of the antiviral factor tetherin[J].Front Immunol,2014,5:655.
[26]Rapicavoli N A,Qu K,Zhang J,Mikhail M,Laberge R-M,Chang H Y.A mammalian pseudogene lncRNAat the interface of inflammation and anti-inflammatory therapeutics[J/OL].Elife,2013,2:e00762.
[27]Liu B,Sun L,Liu Q,Gong C,Yao Y,Lv X,Lin L,Yao H,Su F,Li D,Zeng M,Song E.A cytoplasmic NF-kappaB interacting long noncoding RNA blocks IkappaB phosphorylation and suppresses breast cancer metastasis[J].Cancer Cell,2015,27(3):370-381.
[28]Krawczyk M,Emerson B M.p50-associated COX-2extragenic RNA(PACER)activates COX-2 gene expression by occluding repressive NF-kappaBcomplexes[J/OL].Elife,2014,3:e01776.
[29]Li Z,Chao T-C,Chang K-Y,Lin N,Patil V S,Shimizu C,Head S R,Burns J C,Rana T M.The long noncoding RNA THRIL regulates TNFalpha expression through its interaction with hnRNPL[J].Proc Natl Acad Sci U S A,2014,111(3):1002-1007.
[30]Iiott N E,Heward J A,Roux B,Tsitsiou E,Fenwick P S,Lenzi L,Goodhead I,Hertz-Fowler C,Heger A,Hall N,Donnelly L E,Sims D,Lindsay M A.Long non-coding RNAs and enhancer RNAs regulate the lipopolysaccharide-induced inflammatory response in human monocytes[J].Nat Commun,2014,5:3979.
[31]Lu J,Wu X,Hong M,Tobias P,Han J.A potential suppressive effect of natural antisense IL-1beta RNA on lipopolysaccharide-induced IL-1beta expression[J].JImmunol,2013,190(12):6570-6578.
[32]Mathews M B.Genes for VA-RNA in adenovirus 2[J].Cell,1975,6(2):223-229.
[33]Rossetto C C,Pari G S.Kaposi′s sarcoma-associated herpesvirus noncoding polyadenylated nuclear RNAinteracts with virus-and host cell-encoded proteins and suppresses expression of genes involved in immune modulation[J].J Virol,2011,85(24):13290-13297.
[34]Rossetto C C,Tarrant-Elorza M,Verma S,Purushothaman P,Pari G S.Regulation of viral and cellular gene expression by Kaposi′s sarcoma-associated herpesvirus polyadenylated nuclear RNA[J].J Virol,2013,87(10):5540-5553.
[35]Bidet K,Dadlani D,Garcia-Blanco M A.G3BP1,G3BP2 and CAPRIN1 are required for translation of interferon stimulated mRNAs and are targeted by a dengue virus non-coding RNA[J/OL].PLoS Pathog,2014,10(7):e1004242.
[36]Chang R-Y,Hsu T-W,Chen Y-L,Liu S-F,Tsai Y-J,Lin Y-T,Chen Y-S,Fan Y-H.Japanese encephalitis virus non-coding RNA inhibits activation of interferon by blocking nuclear translocation of interferon regulatory factor 3[J].Vet Microbiol,2013,166(1-2):11-21.
[37]Schuessler A,Funk A,Lazear H M,Cooper D A,Torres S,Daffis S,Jha B K,Kumagai Y,Takeuchi O,Hertzog P,Silverman R,Akira S,Barton D J,Diamond M S,Khromykh A A.West Nile virus noncoding subgenomic RNA contributes to viral evasion of the type I interferon-mediated antiviral response[J].JVirol,2012,86(10):5708-5718.
[38]Mathews M B.Structure,function,and evolution of adenovirus virus-associated RNAs[J].Curr Top Microbiol Immunol,1995,199(Pt 2):173-187.
[39]Nanbo A,Inoue K,Adachi-Takasawa K,Takada K.Epstein-Barr virus RNA confers resistance to interferonalpha-induced apoptosis in Burkitt′s lymphoma[J].Embo J,2002,21(5):954-965.