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Keigo Machida, PhD

TitleAssociate Professor of Molecular Microbiology & Immunology
InstitutionUniversity of Southern California
DepartmentMolecular Microbiology and Immunology
AddressHMR 503C 2011 Zonal Avenue
Health Sciences Campus
Los Angeles CA 90089
Phone+1 323 442 2692
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    Collapse Biography 
    Collapse Education and Training
    University of Tokyo, School of Medicine, Tokyo, JapanPh.D.03/2001Molecular Immunology
    University of Southern California, Los Angeles, CAPost-doc05/2003Molecular Virology
    Howard Hughes Medical Institute, Los Angeles, CAPost-doc08/2005Cancer Biology
    Collapse Awards and Honors
    Japan Society for the Promotion of Science1998  - 2001Scholarship
    Asian Pacific Association for the Study of the Liver (APASL)2007Young Investigator Award
    Research Center for Liver Diseases 2007  - 2007USC Research Center for Liver Diseases Pilot Funding Award
    Liver Meeting of AASLD2008Presidential Poster of Distinction
    American Cancer Society (ACS) 2011  - 2016Research Scholar

    Collapse Overview 
    Collapse Overview
    Hepatitis C Virus (HCV) is a “silent stalker” in infected people, who can develop hepatocellular carcinomas (HCC) decades later. Our short-term goal is to study three areas:

    (i) Host-pathogen interactions to identify the mechanism of HCV-associated HCC

    (ii) Molecular virology of lymphotropic HCV in viral persistence and pathogenesis

    (iii) Cancer biology in liver oncogenesis generated from tumor-initiating cells (TICs: “bad seeds” of treatment-resistant HCC) induced by interactions between environmental factors (alcohol, obesity) and HCV.

    Our lab’s long-term goal is to test new therapeutic modalities for HCC in HCV-infected patients with alcoholism and/or obesity.

    Our experience and resources on TICs have provided a unique opportunity to functionally dissect the alcohol-mediated self-renewal of liver TICs via TLR4 signaling. The p53 tumor suppressor acts as a barrier against stem cell proliferation, and inactivation of p53, or its stabilizing partner NUMB (cell fate decision molecule), leads to expansion of TICs. Although treatment efficacy of HCV has been dramatically improved in recent years, the incidence of HCV-associated HCC keeps rising due to the prevalence of obesity, alcoholism and illicit drug usage in HCV patients. Thus, we will continue our research on the development of new therapeutic modalities for HCC.


    Collapse Research 
    Collapse Research Activities and Funding
    NANOG-positive cancer stem cells in liver oncogenesis induced by alcohol and HCV
    NIH/NIAAA R01AA025204Sep 1, 2017 - May 31, 2022
    Role: Principal Investigator
    Southern California Research Center for ALPD and Cirrhosis
    NIH/NIAAA P50 AA11999Jan 1, 2015 - Dec 31, 2019
    Role: Co-director of Animal Core Facility and Project 3 leader
    Role Description: This center grant supports our central research theme: elucidation of mechanisms by which alcohol sensitizes and primes the liver and pancreas for injury. Animal Core provides the intragastric ethanol infusion model to the Center members.
    Nanog-positive cancer stem cells in and liver oncogenesis by alcohol and HCV
    NIH/NIAAA R01AA018857Sep 15, 2009 - Aug 31, 2014
    Role: Principal Investigator

    Collapse ORNG Applications 
    Collapse Featured Publications
    Collapse Websites
    Collapse Featured Presentations

    Collapse Bibliographic 
    Collapse Publications
    Publications listed below are automatically derived from MEDLINE/PubMed and other sources, which might result in incorrect or missing publications. Researchers can login to make corrections and additions, or contact us for help.
    List All   |   Timeline
    1. Machida K. Pluripotency transcription factors and Metabolic reprogramming of mitochondria in tumor-initiating stem-like cells. Antioxid Redox Signal. 2017 Dec 19. PMID: 29256636.
      View in: PubMed
    2. Liu K, Lee J, Kim JY, Wang L, Tian Y, Chan ST, Cho C, Machida K, Chen D, Ou JJ. Mitophagy Controls the Activities of Tumor Suppressor p53 to Regulate Hepatic Cancer Stem Cells. Mol Cell. 2017 Oct 19; 68(2):281-292.e5. PMID: 29033320.
      View in: PubMed
    3. Baulies A, Montero J, Matías N, Insausti N, Terrones O, Basañez G, Vallejo C, Conde de La Rosa L, Martinez L, Robles D, Morales A, Abian J, Carrascal M, Machida K, Kumar DBU, Tsukamoto H, Kaplowitz N, Garcia-Ruiz C, Fernández-Checa JC. The 2-oxoglutarate carrier promotes liver cancer by sustaining mitochondrial GSH despite cholesterol loading. Redox Biol. 2018 Apr; 14:164-177. PMID: 28942194.
      View in: PubMed
    4. Chen CL, Huang JY, Wang CH, Tahara SM, Zhou L, Kondo Y, Schechter J, Su L, Lai MM, Wakita T, Cosset FL, Jung JU, Machida K. Hepatitis C virus has a genetically determined lymphotropism through co-receptor B7.2. Nat Commun. 2017 Jan 09; 8:13882. PMID: 28067225.
      View in: PubMed
    5. Chen J, Shukla V, Farci P, Andricovich J, Jogunoori W, Kwong LN, Katz LH, Shetty K, Rashid A, Su X, White J, Li L, Wang AY, Blechacz B, Raju GS, Davila M, Nguyen BN, Stroehlein JR, Chen J, Kim SS, Levin H, Machida K, Tsukamoto H, Michaely P, Tzatsos A, Mishra B, Amdur R, Mishra L. Loss of the transforming growth factor-ß effector ß2-Spectrin promotes genomic instability. Hepatology. 2017 Feb; 65(2):678-693. PMID: 28114741.
      View in: PubMed
    6. Machida K. Existence of cancer stem cells in hepatocellular carcinoma: myth or reality? Hepatol Int. 2017 Mar; 11(2):143-147. PMID: 27990610.
      View in: PubMed
    7. Chen J, Yao ZX, Chen JS, Gi YJ, Muñoz NM, Kundra S, Herlong HF, Jeong YS, Goltsov A, Ohshiro K, Mistry NA, Zhang J, Su X, Choufani S, Mitra A, Li S, Mishra B, White J, Rashid A, Wang AY, Javle M, Davila M, Michaely P, Weksberg R, Hofstetter WL, Finegold MJ, Shay JW, Machida K, Tsukamoto H, Mishra L. TGF-ß/ß2-spectrin/CTCF-regulated tumor suppression in human stem cell disorder Beckwith-Wiedemann syndrome. J Clin Invest. 2016 Feb; 126(2):527-42. PMID: 26784546; PMCID: PMC4731168.
    8. Chen CL, Uthaya Kumar DB, Punj V, Xu J, Sher L, Tahara SM, Hess S, Machida K. NANOG Metabolically Reprograms Tumor-Initiating Stem-like Cells through Tumorigenic Changes in Oxidative Phosphorylation and Fatty Acid Metabolism. Cell Metab. 2016 Jan 12; 23(1):206-19. PMID: 26724859; PMCID: PMC4715587 [Available on 01/12/17].
    9. Uthaya Kumar DB, Chen CL, Liu JC, Feldman DE, Sher LS, French S, DiNorcia J, French SW, Naini BV, Junrungsee S, Agopian VG, Zarrinpar A, Machida K. TLR4 Signaling via NANOG Cooperates With STAT3 to Activate Twist1 and Promote Formation of Tumor-Initiating Stem-Like Cells in Livers of Mice. Gastroenterology. 2016 Mar; 150(3):707-19. PMID: 26582088; PMCID: PMC4766021 [Available on 03/01/17].
    10. Siddique HR, Feldman DE, Chen CL, Punj V, Tokumitsu H, Machida K. NUMB phosphorylation destabilizes p53 and promotes self-renewal of tumor-initiating cells by a NANOG-dependent mechanism in liver cancer. Hepatology. 2015 Nov; 62(5):1466-79. PMID: 26174965; PMCID: PMC4618247 [Available on 11/01/16].
    11. He S, Zhao Z, Yang Y, O'Connell D, Zhang X, Oh S, Ma B, Lee JH, Zhang T, Varghese B, Yip J, Dolatshahi Pirooz S, Li M, Zhang Y, Li GM, Ellen Martin S, Machida K, Liang C. Truncating mutation in the autophagy gene UVRAG confers oncogenic properties and chemosensitivity in colorectal cancers. Nat Commun. 2015 Aug 03; 6:7839. PMID: 26234763; PMCID: PMC4526116.
    12. Feng T, Dzieran J, Gu X, Marhenke S, Vogel A, Machida K, Weiss TS, Ruemmele P, Kollmar O, Hoffmann P, Grässer F, Allgayer H, Fabian J, Weng HL, Teufel A, Maass T, Meyer C, Lehmann U, Zhu C, Mertens PR, Gao CF, Dooley S, Meindl-Beinker NM. Smad7 regulates compensatory hepatocyte proliferation in damaged mouse liver and positively relates to better clinical outcome in human hepatocellular carcinoma. Clin Sci (Lond). 2015 Jun; 128(11):761-74. PMID: 25602745.
      View in: PubMed
    13. Machida K, Feldman DE, Tsukamoto H. TLR4-dependent tumor-initiating stem cell-like cells (TICs) in alcohol-associated hepatocellular carcinogenesis. Adv Exp Med Biol. 2015; 815:131-44. PMID: 25427905.
      View in: PubMed
    14. Lazaro R, Wu R, Lee S, Zhu NL, Chen CL, French SW, Xu J, Machida K, Tsukamoto H. Osteopontin deficiency does not prevent but promotes alcoholic neutrophilic hepatitis in mice. Hepatology. 2015 Jan; 61(1):129-40. PMID: 25132354; PMCID: PMC4280361.
    15. Takemoto K, Hatano E, Iwaisako K, Takeiri M, Noma N, Ohmae S, Toriguchi K, Tanabe K, Tanaka H, Seo S, Taura K, Machida K, Takeda N, Saji S, Uemoto S, Asagiri M. Necrostatin-1 protects against reactive oxygen species (ROS)-induced hepatotoxicity in acetaminophen-induced acute liver failure. FEBS Open Bio. 2014; 4:777-87. PMID: 25349782; PMCID: PMC4208088.
    16. Chen CL, Tsukamoto H, Machida K. Oncogenic signaling pathways and origins of tumor-initiating stem-like cells of hepatocellular carcinomas induced by hepatitis C virus, alcohol and/or obesity. Hepatol Int. 2014 Jul; 8(3):330-8. PMID: 26202636.
      View in: PubMed
    17. Kondo Y, Ninomiya M, Kimura O, Machida K, Funayama R, Nagashima T, Kobayashi K, Kakazu E, Kato T, Nakayama K, Lai MM, Shimosegawa T. HCV infection enhances Th17 commitment, which could affect the pathogenesis of autoimmune diseases. PLoS One. 2014; 9(6):e98521. PMID: 24905921; PMCID: PMC4048196.
    18. Tsukamoto H, Mishra L, Machida K. Alcohol, TLR4-TGF-ß antagonism, and liver cancer. Hepatol Int. 2014 Sep; 8 Suppl 2:408-12. PMID: 26201318; PMCID: PMC4900455.
    19. Chen CL, Tsukamoto H, Liu JC, Kashiwabara C, Feldman D, Sher L, Dooley S, French SW, Mishra L, Petrovic L, Jeong JH, Machida K. Reciprocal regulation by TLR4 and TGF-ß in tumor-initiating stem-like cells. J Clin Invest. 2013 Jul; 123(7):2832-49. PMID: 23921128; PMCID: PMC3696549.
    20. Machida K. Tumor-initiating stem-like cells and drug resistance: carcinogenesis through Toll-like receptors, environmental factors, and virus. Drug Deliv Transl Res. 2013 Apr; 3(2):152-64. PMID: 25787983.
      View in: PubMed
    21. Feldman DE, Chen C, Punj V, Machida K. The TBC1D15 oncoprotein controls stem cell self-renewal through destabilization of the Numb-p53 complex. PLoS One. 2013; 8(2):e57312. PMID: 23468968; PMCID: PMC3584131.
    22. Kondo Y, Uno K, Machida K, Terajima M. PPARs and Liver Disease. PPAR Res. 2013; 2013:896412. PMID: 23476632; PMCID: PMC3576795.
    23. Hong B, Lee SH, Song XT, Jones L, Machida K, Huang XF, Chen SY. A super TLR agonist to improve efficacy of dendritic cell vaccine in induction of anti-HCV immunity. PLoS One. 2012; 7(11):e48614. PMID: 23144910; PMCID: PMC3492467.
    24. Liu Z, Tian Y, Machida K, Lai MM, Luo G, Foung SK, Ou JH. Transient activation of the PI3K-AKT pathway by hepatitis C virus to enhance viral entry. J Biol Chem. 2012 Dec 07; 287(50):41922-30. PMID: 23095753; PMCID: PMC3516739.
    25. Liu HM, Aizaki H, Machida K, Ou JH, Lai MM. Hepatitis C virus translation preferentially depends on active RNA replication. PLoS One. 2012; 7(8):e43600. PMID: 22937067; PMCID: PMC3427374.
    26. von Montfort C, Matias N, Fernandez A, Fucho R, Conde de la Rosa L, Martinez-Chantar ML, Mato JM, Machida K, Tsukamoto H, Murphy MP, Mansouri A, Kaplowitz N, Garcia-Ruiz C, Fernandez-Checa JC. Mitochondrial GSH determines the toxic or therapeutic potential of superoxide scavenging in steatohepatitis. J Hepatol. 2012 Oct; 57(4):852-9. PMID: 22687340; PMCID: PMC3445734.
    27. Sir D, Kuo CF, Tian Y, Liu HM, Huang EJ, Jung JU, Machida K, Ou JH. Replication of hepatitis C virus RNA on autophagosomal membranes. J Biol Chem. 2012 May 25; 287(22):18036-43. PMID: 22496373; PMCID: PMC3365724.
    28. Ueno A, Lazaro R, Wang PY, Higashiyama R, Machida K, Tsukamoto H. Mouse intragastric infusion (iG) model. Nat Protoc. 2012 Mar 29; 7(4):771-81. PMID: 22461066; PMCID: PMC4197794.
    29. Tsukamoto H, Zhu NL, Wang J, Asahina K, Machida K. Morphogens and hepatic stellate cell fate regulation in chronic liver disease. J Gastroenterol Hepatol. 2012 Mar; 27 Suppl 2:94-8. PMID: 22320925; PMCID: PMC3337168.
    30. Machida K, Chen CL, Liu JC, Kashiwabara C, Feldman D, French SW, Sher L, Hyeongnam JJ, Tsukamoto H. Cancer stem cells generated by alcohol, diabetes, and hepatitis C virus. J Gastroenterol Hepatol. 2012 Mar; 27 Suppl 2:19-22. PMID: 22320911; PMCID: PMC3306127.
    31. Feldman DE, Chen C, Punj V, Tsukamoto H, Machida K. Pluripotency factor-mediated expression of the leptin receptor (OB-R) links obesity to oncogenesis through tumor-initiating stem cells. Proc Natl Acad Sci U S A. 2012 Jan 17; 109(3):829-34. PMID: 22207628; PMCID: PMC3271911.
    32. Inn KS, Lee SH, Rathbun JY, Wong LY, Toth Z, Machida K, Ou JH, Jung JU. Inhibition of RIG-I-mediated signaling by Kaposi's sarcoma-associated herpesvirus-encoded deubiquitinase ORF64. J Virol. 2011 Oct; 85(20):10899-904. PMID: 21835791; PMCID: PMC3187500.
    33. Tsukiyama-Kohara K, Sekiguchi S, Kasama Y, Salem NE, Machida K, Kohara M. Hepatitis C virus-related lymphomagenesis in a mouse model. ISRN Hematol. 2011; 2011:167501. PMID: 22084693; PMCID: PMC3195281.
    34. Szabo G, Billiar TR, Machida K, Crispe IN, Seki E. Toll-like receptor signaling in liver diseases. Gastroenterol Res Pract. 2010; 2010:971270. PMID: 21789039; PMCID: PMC3123973.
    35. Machida K. TLRs, Alcohol, HCV, and Tumorigenesis. Gastroenterol Res Pract. 2010; 2010:518674. PMID: 21331379; PMCID: PMC3038599.
    36. Zhong S, Machida K, Tsukamoto H, Johnson DL. Alcohol induces RNA polymerase III-dependent transcription through c-Jun by co-regulating TATA-binding protein (TBP) and Brf1 expression. J Biol Chem. 2011 Jan 28; 286(4):2393-401. PMID: 21106530; PMCID: PMC3024733.
    37. Machida K, McNamara G, Cheng KT, Huang J, Wang CH, Comai L, Ou JH, Lai MM. Hepatitis C virus inhibits DNA damage repair through reactive oxygen and nitrogen species and by interfering with the ATM-NBS1/Mre11/Rad50 DNA repair pathway in monocytes and hepatocytes. J Immunol. 2010 Dec 01; 185(11):6985-98. PMID: 20974981; PMCID: PMC3101474.
    38. Yeligar SM, Machida K, Kalra VK. Ethanol-induced HO-1 and NQO1 are differentially regulated by HIF-1alpha and Nrf2 to attenuate inflammatory cytokine expression. J Biol Chem. 2010 Nov 12; 285(46):35359-73. PMID: 20833713; PMCID: PMC2975160.
    39. Lai CK, Jeng KS, Machida K, Lai MM. Hepatitis C virus egress and release depend on endosomal trafficking of core protein. J Virol. 2010 Nov; 84(21):11590-8. PMID: 20739534; PMCID: PMC2953145.
    40. Kondo Y, Ueno Y, Kakazu E, Kobayashi K, Shiina M, Tamai K, Machida K, Inoue J, Wakui Y, Fukushima K, Obara N, Kimura O, Shimosegawa T. Lymphotropic HCV strain can infect human primary naïve CD4+ cells and affect their proliferation and IFN-? secretion activity. J Gastroenterol. 2011 Feb; 46(2):232-41. PMID: 20714907; PMCID: PMC4197795.
    41. Machida K, Tsukamoto H, Liu JC, Han YP, Govindarajan S, Lai MM, Akira S, Ou JH. c-Jun mediates hepatitis C virus hepatocarcinogenesis through signal transducer and activator of transcription 3 and nitric oxide-dependent impairment of oxidative DNA repair. Hepatology. 2010 Aug; 52(2):480-92. PMID: 20683948; PMCID: PMC3107125.
    42. Chen YC, Su WC, Huang JY, Chao TC, Jeng KS, Machida K, Lai MM. Polo-like kinase 1 is involved in hepatitis C virus replication by hyperphosphorylating NS5A. J Virol. 2010 Aug; 84(16):7983-93. PMID: 20534861; PMCID: PMC2916529.
    43. Szabo G, Wands JR, Eken A, Osna NA, Weinman SA, Machida K, Joe Wang H. Alcohol and hepatitis C virus--interactions in immune dysfunctions and liver damage. Alcohol Clin Exp Res. 2010 Oct; 34(10):1675-86. PMID: 20608905; PMCID: PMC3253556.
    44. Yeligar SM, Machida K, Tsukamoto H, Kalra VK. Ethanol augments RANTES/CCL5 expression in rat liver sinusoidal endothelial cells and human endothelial cells via activation of NF-kappa B, HIF-1 alpha, and AP-1. J Immunol. 2009 Nov 01; 183(9):5964-76. PMID: 19828633; PMCID: PMC3633447.
    45. Machida K, Liu JC, McNamara G, Levine A, Duan L, Lai MM. Hepatitis C virus causes uncoupling of mitotic checkpoint and chromosomal polyploidy through the Rb pathway. J Virol. 2009 Dec; 83(23):12590-600. PMID: 19793824; PMCID: PMC2786750.
    46. Tsukamoto H, Machida K, Dynnyk A, Mkrtchyan H. "Second hit" models of alcoholic liver disease. Semin Liver Dis. 2009 May; 29(2):178-87. PMID: 19387917; PMCID: PMC4197797.
    47. Machida K, Tsukiyama-Kohara K, Sekiguch S, Seike E, Tóne S, Hayashi Y, Tobita Y, Kasama Y, Shimizu M, Takahashi H, Taya C, Yonekawa H, Tanaka N, Kohara M. Hepatitis C virus and disrupted interferon signaling promote lymphoproliferation via type II CD95 and interleukins. Gastroenterology. 2009 Jul; 137(1):285-96, 296.e1-11. PMID: 19362089; PMCID: PMC4197798.
    48. Kondo Y, Machida K, Liu HM, Ueno Y, Kobayashi K, Wakita T, Shimosegawa T, Lai MM. Hepatitis C virus infection of T cells inhibits proliferation and enhances fas-mediated apoptosis by down-regulating the expression of CD44 splicing variant 6. J Infect Dis. 2009 Mar 01; 199(5):726-36. PMID: 19199548; PMCID: PMC4174598.
    49. Liu HM, Aizaki H, Choi KS, Machida K, Ou JJ, Lai MM. SYNCRIP (synaptotagmin-binding, cytoplasmic RNA-interacting protein) is a host factor involved in hepatitis C virus RNA replication. Virology. 2009 Apr 10; 386(2):249-56. PMID: 19232660; PMCID: PMC3099193.
    50. Machida K, Tsukamoto H, Mkrtchyan H, Duan L, Dynnyk A, Liu HM, Asahina K, Govindarajan S, Ray R, Ou JH, Seki E, Deshaies R, Miyake K, Lai MM. Toll-like receptor 4 mediates synergism between alcohol and HCV in hepatic oncogenesis involving stem cell marker Nanog. Proc Natl Acad Sci U S A. 2009 Feb 03; 106(5):1548-53. PMID: 19171902; PMCID: PMC2635765.
    51. Lai CK, Jeng KS, Machida K, Lai MM. Association of hepatitis C virus replication complexes with microtubules and actin filaments is dependent on the interaction of NS3 and NS5A. J Virol. 2008 Sep; 82(17):8838-48. PMID: 18562541; PMCID: PMC2519648.
    52. Machida K, Kondo Y, Huang JY, Chen YC, Cheng KT, Keck Z, Foung S, Dubuisson J, Sung VM, Lai MM. Hepatitis C virus (HCV)-induced immunoglobulin hypermutation reduces the affinity and neutralizing activities of antibodies against HCV envelope protein. J Virol. 2008 Jul; 82(13):6711-20. PMID: 18417597; PMCID: PMC2447061.
    53. Lai CK, Jeng KS, Machida K, Cheng YS, Lai MM. Hepatitis C virus NS3/4A protein interacts with ATM, impairs DNA repair and enhances sensitivity to ionizing radiation. Virology. 2008 Jan 20; 370(2):295-309. PMID: 17931678.
      View in: PubMed
    54. Kondo Y, Sung VM, Machida K, Liu M, Lai MM. Hepatitis C virus infects T cells and affects interferon-gamma signaling in T cell lines. Virology. 2007 Apr 25; 361(1):161-73. PMID: 17175001.
      View in: PubMed
    55. Kawamura H, Govindarajan S, Aswad F, Machida K, Lai MM, Sung VM, Dennert G. HCV core expression in hepatocytes protects against autoimmune liver injury and promotes liver regeneration in mice. Hepatology. 2006 Oct; 44(4):936-44. PMID: 17006910.
      View in: PubMed
    56. Machida K, Cheng KT, Lai CK, Jeng KS, Sung VM, Lai MM. Hepatitis C virus triggers mitochondrial permeability transition with production of reactive oxygen species, leading to DNA damage and STAT3 activation. J Virol. 2006 Jul; 80(14):7199-207. PMID: 16809325; PMCID: PMC1489016.
    57. Machida K, Cheng KT, Sung VM, Levine AM, Foung S, Lai MM. Hepatitis C virus induces toll-like receptor 4 expression, leading to enhanced production of beta interferon and interleukin-6. J Virol. 2006 Jan; 80(2):866-74. PMID: 16378988; PMCID: PMC1346849.
    58. Machida K, Cheng KT, Pavio N, Sung VM, Lai MM. Hepatitis C virus E2-CD81 interaction induces hypermutation of the immunoglobulin gene in B cells. J Virol. 2005 Jul; 79(13):8079-89. PMID: 15956553; PMCID: PMC1143751.
    59. Machida K, Cheng KT, Sung VM, Lee KJ, Levine AM, Lai MM. Hepatitis C virus infection activates the immunologic (type II) isoform of nitric oxide synthase and thereby enhances DNA damage and mutations of cellular genes. J Virol. 2004 Aug; 78(16):8835-43. PMID: 15280491; PMCID: PMC479064.
    60. Machida K, Cheng KT, Sung VM, Shimodaira S, Lindsay KL, Levine AM, Lai MY, Lai MM. Hepatitis C virus induces a mutator phenotype: enhanced mutations of immunoglobulin and protooncogenes. Proc Natl Acad Sci U S A. 2004 Mar 23; 101(12):4262-7. PMID: 14999097; PMCID: PMC384729.
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