miR-122
mir-122 microRNA precursor | |
---|---|
Identifiers | |
Symbol | mir-122 |
Rfam | RF00684 |
miRBase | MI0000442 |
miRBase family | MIPF0000095 |
Other data | |
RNA type | Gene; miRNA |
Domain(s) | Eukaryota |
GO | GO:0035195 GO:0035068 |
SO | SO:0001244 |
PDB structures | PDBe |
miR-122 is a miRNA that is conserved among vertebrate species. miR-122 is not present in invertebrates, and no close paralogs of miR-122 have been detected.[1] miR-122 is highly expressed in the liver, where it has been implicated as a regulator of fatty-acid metabolism in mouse studies. Reduced miR-122 levels are associated with hepatocellular carcinoma. miR-122 also plays an important positive role in the regulation of hepatitis C virus replication.
Expression and regulation
miR-122 was originally identified by cloning of tissue-specific microRNAs in mouse, where its expression is restricted to the liver.[2] The liver-specific expression of miR-122 is conserved in zebrafish.[3] miR-122 expression increases during embryogenesis until it constitutes 72% of total miRNA in adult human liver, making it one of the most highly expressed miRNAs in any tissue.[4] In humans, miR-122 is encoded at a single genomic locus in chromosome 18. The primary miR-122 transcript (pri-miR-122) is a long non-coding RNA. Transcription is regulated by HNF4α.[5] The miR-122 hairpin precursor consensus shown here is predicted based on base pairing and cross-species conservation. The mature sequence is excised from the 5' arm of the hairpin.[2][6]
There is evidence that miR-122 is regulated by Rev-ErbA alpha which is involved in circadian gene expression, suggesting that miR-122 is a circadian metabolic regulator. miR-122 regulates the expression of several mRNA molecules that are important in the circadian cycle, such as PPARβ/δ.[7] Mature miR-122 is subject to modification by the poly(A) polymerase GLD-2, which adds a single adenosine to the miRNA 3' end. This results in an increase in miR-122 stability.[8]
Targets
miR-122 regulates the synthesis of the protein CAT-1 by binding to sites in the mRNA 3'UTR such that translation is repressed and the mRNA is targeted to P bodies. This repression can be relieved by the protein HuR, which is released from the nucleus under conditions of cell stress and binds to the CAT-1 3'UTR. The HuR interaction leads to release of the mRNA from the P bodies and resumption of active translation.[9]
A number of other miR-122 targets, including CD320, AldoA and BCKDK, have been identified by microarray analysis of changes in mRNA expression in the liver of mice treated with miR-122 inhibitors.[10][11][12] The overall effect of miR-122 inhibition is to reduce the plasma cholesterol level, although the pathways involved in this regulation have not been fully elucidated. miR-122 also regulates systemic iron homeostasis via the target mRNAs Hjv and Hfe.[13] miR-122 inhibition in mice or primates does not result in any detectable liver toxicity.[14]
Role in cancer
miR-122 levels are frequently reduced in hepatocellular carcinoma (HCC) compared to normal liver, and low miR-122 levels correlate with poor prognosis.[15][16] Overexpression of miR-122 reduces tumorigenic properties in HCC cell lines, suggesting that it functions as a tumor suppressor gene, and increases the response of cells to the chemotherapeutic drugs sorafenib and doxorubicin.[17][18] Several miR-122 target genes have been implicated in tumorigenesis, including ADAM10, IGF1R, CCNG1 and ADAM17.[17][18][19]
Innate Immunity
Recent studies demonstrated that miR-122 may directly regulate different aspects of the interferons (IFNs) signaling pathway[20][21] to enhanced induction of anti-viral genes and inhibition of various virus.[21][22][23][24][25][26][27][28][29][30] Moreover, miR-122 have been shown to target various genes,[31][32][29][33][28] resulting in enhancement of IFN signaling and subsequent antiviral innate immunity.[31][34] Interferons (IFNs, includes type I and III interferon) treatment leads to a significant reduction in the expression of the liver-specific miR-122.[21][35][36][37][28] HepG2 cells with overexpressed microRNA-122 mount an effective antiviral interferon response and innate immune response to hepatitis C virus (HCV), other RNA viruses and viral mimetics (e.g. poly(I:C)).[22]
Regulation of HCV
Recent studies have shown that replication of hepatitis C virus (HCV) is dependent on miR-122 expression.[38] miR-122 regulates HCV by binding directly to two adjacent sites close to the 5' end of HCV RNA.[39] Although these experiments were conducted using genotype 1a and 1b HCV RNA, the miR-122 binding sites are highly conserved across different genotypes, and miR-122 is also required for replication of infectious type 2a HCV.[40] As miRNAs generally function to repress gene expression by binding to 3'UTR sites, this positive regulation of viral replication via a 5'UTR represents a novel function for miR-122. The mechanism of regulation is not yet clear. miR-122 stimulates translation of HCV RNA, but not to a sufficient extent to explain its effects on viral replication, indicating that a second stage of the viral replication cycle must also be regulated. [41][42] HCV RNA synthesis is not affected by miR-122, suggesting that regulation of other processes such as RNA stability may occur.[43][44] The extent to which the miRNA-induced silencing complex (miRISC) is involved in this regulation has not been fully determined. The Argonaute proteins (Ago1–4), which are essential for miRNA-directed repression, appear to be necessary for miR-122 to regulate HCV,[45] although miR-122 overexpression may overcome this requirement.[46] The crystal structure of Ago2:miR-122 bound to the miR-122 binding site at the 5'-end of the HCV genome, in combination with functional experiments, suggests that the viral RNA has evolved to maximize protection from cytoplasmic exoribonucleases by altering the molecular behavior of Ago2.[47] Another miRISC component, the DEAD-box RNA helicase DDX6, does not play a role in miR-122-facilitated HCV replication.[48]
The existing HCV therapy of PEG-IFNα plus ribavirin is poorly tolerated and frequently ineffective,[49][26] so there is an urgent need for new drugs, and miR-122 inhibitors are an attractive possibility. The association between low miR-122 levels and hepatocellular carcinoma suggests that caution will be necessary when testing miR-122 inhibitors, and that long-term treatment might be undesirable. However, miR-122 is a promising target as it can be very selectively and effectively inhibited with antisense oligonucleotides, and as it is a conserved host factor it is hoped that the virus would not be able to acquire resistance mutations to an anti-miR-122 therapeutic. Moreover, engineering HepG2 cells to express miR-122 (HepG2-HFL cell, HepG2 cells expressing miR-122) mount an effective antiviral interferon-lambda (IFNλ) based innate immune response to hepatitis C virus (HCV) infection.[22][25] HepG2 cells (stably expressing miR-122) produced a more robust IFN Response (type I and type III interferons) when challenged with other RNA viruses [ IAV-ΔNS1 and SeV ] and viral mimetics than Huh-7 and Huh-7.5 cells. HCV Induces an IFN-λ (IL28 and IL29), ISG, and Cytokine Response in these HepG2 cells with stably expressing miR-122.[22][23][24][31][34]
Inhibitor miravirsen
As of 2017, Santaris Pharma was developing miravirsen, a locked nucleic acid-based antisense oligonucleotide that inhibits miR-122, as a potential treatment for HepC.[50]
Use as a biomarker
miR-122 has recently been explored as a potential biomarker for various hepatic conditions. A change in levels of miR-122 in the blood has been confirmed as an indicator for viral-, alcohol- and chemical-induced liver injury[51][52][53] as well as Transplant rejection after Liver transplantation.[54][55] This change is noted before increased amino-transferase activity, making it an early indicator of liver disease and hepatocellular injury of liver grafts prior to liver transplantation.[54][56]
There is a great deal of research into the use of miR-122 as a biomarker for hepatitis C. While some studies dispute its efficacy for diagnosing Hep C,[57] other research indicates that it may be useful in diagnosing specific forms of hepatitis.[58] In addition, decreased levels of miR-122 in liver biopsies have been linked to a strain of hepatitis C that is resistant to interferon therapy.[59]
miR-122 has also been suggested as a biomarker for hepatectomy-induced liver injury in patients with hepatocellular carcinoma.[60]
It is important to note that detection of miR-122 and other microRNAs in Body fluid like blood can be interfered by heparin contaminated. The commonly used anticoagulant heparin profoundly inhibits the by reverse transcription polymerase chain reaction (RT-PCR) used for microRNA quantification.[61][62]
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Further reading
- Zeng C, Wang R, Li D, Lin XJ, Wei QK, Yuan Y, et al. (November 2010). "A novel GSK-3 beta-C/EBP alpha-miR-122-insulin-like growth factor 1 receptor regulatory circuitry in human hepatocellular carcinoma". Hepatology. 52 (5): 1702–1712. doi:10.1002/hep.23875. PMID 21038412. S2CID 25688272.
- Shea CM, Tzertzinis G (October 2010). "Controlled expression of functional miR-122 with a ligand inducible expression system". BMC Biotechnology. 10 (1): 76. doi:10.1186/1472-6750-10-76. PMC 2976731. PMID 20961424.
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: CS1 maint: unflagged free DOI (link) - Stilling G, Sun Z, Zhang S, Jin L, Righi A, Kovācs G, et al. (August 2010). "MicroRNA expression in ACTH-producing pituitary tumors: up-regulation of microRNA-122 and -493 in pituitary carcinomas". Endocrine. 38 (1): 67–75. doi:10.1007/s12020-010-9346-0. PMID 20960104. S2CID 207361604.
- Zhang Y, Jia Y, Zheng R, Guo Y, Wang Y, Guo H, et al. (December 2010). "Plasma microRNA-122 as a biomarker for viral-, alcohol-, and chemical-related hepatic diseases". Clinical Chemistry. 56 (12): 1830–1838. doi:10.1373/clinchem.2010.147850. PMID 20930130.
- Xu H, He JH, Xiao ZD, Zhang QQ, Chen YQ, Zhou H, Qu LH (October 2010). "Liver-enriched transcription factors regulate microRNA-122 that targets CUTL1 during liver development". Hepatology. 52 (4): 1431–1442. doi:10.1002/hep.23818. PMID 20842632. S2CID 29273466.
- Burchard J, Zhang C, Liu AM, Poon RT, Lee NP, Wong KF, et al. (August 2010). "microRNA-122 as a regulator of mitochondrial metabolic gene network in hepatocellular carcinoma". Molecular Systems Biology. 6 (1): 402. doi:10.1038/msb.2010.58. PMC 2950084. PMID 20739924.
- Qiu L, Fan H, Jin W, Zhao B, Wang Y, Ju Y, et al. (August 2010). "miR-122-induced down-regulation of HO-1 negatively affects miR-122-mediated suppression of HBV". Biochemical and Biophysical Research Communications. 398 (4): 771–777. doi:10.1016/j.bbrc.2010.07.021. PMID 20633528.
- Lin LT, Noyce RS, Pham TN, Wilson JA, Sisson GR, Michalak TI, et al. (September 2010). "Replication of subgenomic hepatitis C virus replicons in mouse fibroblasts is facilitated by deletion of interferon regulatory factor 3 and expression of liver-specific microRNA 122". Journal of Virology. 84 (18): 9170–9180. doi:10.1128/JVI.00559-10. PMC 2937658. PMID 20592082.
- Kojima S, Gatfield D, Esau CC, Green CB (June 2010). Yamazaki S (ed.). "MicroRNA-122 modulates the rhythmic expression profile of the circadian deadenylase Nocturnin in mouse liver". PLOS ONE. 5 (6): e11264. Bibcode:2010PLoSO...511264K. doi:10.1371/journal.pone.0011264. PMC 2889834. PMID 20582318.
- Qian J, Zhai A, Kao W, Li Y, Song W, Fu Y, et al. (August 2010). "Modulation of miR-122 on persistently Borna disease virus infected human oligodendroglial cells". Antiviral Research. 87 (2): 249–256. doi:10.1016/j.antiviral.2010.05.011. PMID 20561966.
- Young DD, Connelly CM, Grohmann C, Deiters A (June 2010). "Small molecule modifiers of microRNA miR-122 function for the treatment of hepatitis C virus infection and hepatocellular carcinoma". Journal of the American Chemical Society. 132 (23): 7976–7981. doi:10.1021/ja910275u. PMID 20527935.
- Chang Y, He XX, Li PY, Lin JS (April 2010). "[MiR-122 regulates the expression of PEG10 in hepatoma cell lines]". Zhonghua Gan Zang Bing Za Zhi = Zhonghua Ganzangbing Zazhi = Chinese Journal of Hepatology. 18 (4): 288–291. doi:10.3760/cma.j.issn.1007-3418.2010.04.013. PMID 20460050.
- Lee TC, Lin YL, Liao JT, Su CM, Lin CC, Lin WP, Liao CL (June 2010). "Utilizing liver-specific microRNA-122 to modulate replication of dengue virus replicon". Biochemical and Biophysical Research Communications. 396 (3): 596–601. doi:10.1016/j.bbrc.2010.04.080. PMID 20412785.
- Pfeffer S, Baumert TF (April 2010). Kowdley K, McCaughan G, Trautwein C (eds.). "Antagonizing microRNA-122 and treatment of hepatitis C virus infection". Hepatology. 51 (4): 1461–1463. doi:10.1002/hep.23573. PMID 20373371. S2CID 2669614.
- Branch AD, Rice CM (January 2010). "Antisense gets a grip on miR-122 in chimpanzees". Science Translational Medicine. 2 (13): 13ps1. doi:10.1126/scitranslmed.3000605. PMID 20371461. S2CID 206675938.
- Norman KL, Sarnow P (March 2010). "Hepatitis C virus' Achilles' heel--dependence on liver-specific microRNA miR-122". Cell Research. 20 (3): 247–249. doi:10.1038/cr.2010.28. PMID 20190773. S2CID 8304279.
- Ma L, Liu J, Shen J, Liu L, Wu J, Li W, et al. (April 2010). "Expression of miR-122 mediated by adenoviral vector induces apoptosis and cell cycle arrest of cancer cells". Cancer Biology & Therapy. 9 (7): 554–561. doi:10.4161/cbt.9.7.11267. PMID 20150764. S2CID 24824980.
- Haussecker D, Kay MA (February 2010). "miR-122 continues to blaze the trail for microRNA therapeutics". Molecular Therapy. 18 (2): 240–242. doi:10.1038/mt.2009.313. PMC 2839286. PMID 20125164.
- Iliopoulos D, Drosatos K, Hiyama Y, Goldberg IJ, Zannis VI (June 2010). "MicroRNA-370 controls the expression of microRNA-122 and Cpt1alpha and affects lipid metabolism". Journal of Lipid Research. 51 (6): 1513–1523. doi:10.1194/jlr.M004812. PMC 3035515. PMID 20124555.
{{cite journal}}
: CS1 maint: unflagged free DOI (link) - Wu X, Wu S, Tong L, Luan T, Lin L, Lu S, et al. (2009). "miR-122 affects the viability and apoptosis of hepatocellular carcinoma cells". Scandinavian Journal of Gastroenterology. 44 (11): 1332–1339. doi:10.3109/00365520903215305. PMID 19891584. S2CID 6899554.
- Huang Z, Liu C (April 2009). "[Construction and identification of the human liver-specific miR-122 expression vector]". Sheng Wu Gong Cheng Xue Bao = Chinese Journal of Biotechnology. 25 (4): 587–590. PMID 19637636.
- Zhang R, Wang L, Yu GR, Zhang X, Yao LB, Yang AG (October 2009). "MicroRNA-122 might be a double-edged sword in hepatocellular carcinoma". Hepatology. 50 (4): 1322–1323. doi:10.1002/hep.23108. PMID 19591123. S2CID 34673179.
- Díaz-Toledano R, Ariza-Mateos A, Birk A, Martínez-García B, Gómez J (September 2009). "In vitro characterization of a miR-122-sensitive double-helical switch element in the 5' region of hepatitis C virus RNA". Nucleic Acids Research. 37 (16): 5498–5510. doi:10.1093/nar/gkp553. PMC 2760801. PMID 19578061.
- Jopling CL (December 2008). "Regulation of hepatitis C virus by microRNA-122". Biochemical Society Transactions. 36 (Pt 6): 1220–1223. doi:10.1042/BST0361220. PMID 19021529.
- Lin CJ, Gong HY, Tseng HC, Wang WL, Wu JL (October 2008). "miR-122 targets an anti-apoptotic gene, Bcl-w, in human hepatocellular carcinoma cell lines" (PDF). Biochemical and Biophysical Research Communications. 375 (3): 315–320. doi:10.1016/j.bbrc.2008.07.154. PMID 18692484.
- Chang J, Guo JT, Jiang D, Guo H, Taylor JM, Block TM (August 2008). "Liver-specific microRNA miR-122 enhances the replication of hepatitis C virus in nonhepatic cells". Journal of Virology. 82 (16): 8215–8223. doi:10.1128/JVI.02575-07. PMC 2519557. PMID 18550664.
- Girard M, Jacquemin E, Munnich A, Lyonnet S, Henrion-Caude A (April 2008). "miR-122, a paradigm for the role of microRNAs in the liver". Journal of Hepatology. 48 (4): 648–656. doi:10.1016/j.jhep.2008.01.019. PMID 18291553.
- Fabani MM, Gait MJ (February 2008). "miR-122 targeting with LNA/2'-O-methyl oligonucleotide mixmers, peptide nucleic acids (PNA), and PNA-peptide conjugates". RNA. 14 (2): 336–346. doi:10.1261/rna.844108. PMC 2212241. PMID 18073344.
- Jopling CL, Norman KL, Sarnow P (2006). "Positive and negative modulation of viral and cellular mRNAs by liver-specific microRNA miR-122". Cold Spring Harbor Symposia on Quantitative Biology. 71: 369–376. doi:10.1101/sqb.2006.71.022. PMID 17381319. S2CID 23102623.
- Conrad KD, Giering F, Erfurth C, Neumann A, Fehr C, Meister G, Niepmann M (2013). "MicroRNA-122 dependent binding of Ago2 protein to hepatitis C virus RNA is associated with enhanced RNA stability and translation stimulation". PLOS ONE. 8 (2): e56272. Bibcode:2013PLoSO...856272C. doi:10.1371/journal.pone.0056272. PMC 3566042. PMID 23405269.
- Mortimer SA, Doudna JA (April 2013). "Unconventional miR-122 binding stabilizes the HCV genome by forming a trimolecular RNA structure". Nucleic Acids Research. 41 (7): 4230–4240. doi:10.1093/nar/gkt075. PMC 3627571. PMID 23416544.
External links