메뉴 건너뛰기




Volumn 34, Issue 19, 2015, Pages 2483-2492

Polyoma small T antigen triggers cell death via mitotic catastrophe

Author keywords

[No Author keywords available]

Indexed keywords

PHOSPHOPROTEIN PHOSPHATASE 2; PROTEIN P53; VIRUS T ANTIGEN;

EID: 84940275909     PISSN: 09509232     EISSN: 14765594     Source Type: Journal    
DOI: 10.1038/onc.2014.192     Document Type: Article
Times cited : (10)

References (52)
  • 1
    • 67349257915 scopus 로고    scopus 로고
    • Cellular transformation by Simian Virus 40 and Murine Polyoma Virus T antigens
    • Cheng J, DeCaprio JA, Fluck MM, Schaffhausen BS. Cellular transformation by Simian Virus 40 and Murine Polyoma Virus T antigens. Semin Cancer Biol 2009; 19: 218-228.
    • (2009) Semin Cancer Biol , vol.19 , pp. 218-228
    • Cheng, J.1    Decaprio, J.A.2    Fluck, M.M.3    Schaffhausen, B.S.4
  • 2
    • 0018348655 scopus 로고
    • Tantigen is bound to a host protein in SV40- transformed cells
    • Lane DP, Crawford LV. Tantigen is bound to a host protein in SV40- transformed cells. Nature 1979; 278: 261-263.
    • (1979) Nature , vol.278 , pp. 261-263
    • Lane, D.P.1    Crawford, L.V.2
  • 3
    • 0018760324 scopus 로고
    • Characterization of a 54K dalton cellular SV40 tumor antigen present in SV40-transformed cells and uninfected embryonal carcinoma cells
    • Linzer DI, Levine AJ. Characterization of a 54K dalton cellular SV40 tumor antigen present in SV40-transformed cells and uninfected embryonal carcinoma cells. Cell 1979; 17: 43-52.
    • (1979) Cell , vol.17 , pp. 43-52
    • Linzer, D.I.1    Levine, A.J.2
  • 4
    • 0021828496 scopus 로고
    • Association of phosphatidylinositol kinase activity with polyoma middle-T competent for transformation
    • Whitman M, Kaplan DR, Schaffhausen B, Cantley L, Roberts TM. Association of phosphatidylinositol kinase activity with polyoma middle-T competent for transformation. Nature 1985; 315: 239-242.
    • (1985) Nature , vol.315 , pp. 239-242
    • Whitman, M.1    Kaplan, D.R.2    Schaffhausen, B.3    Cantley, L.4    Roberts, T.M.5
  • 5
    • 0025103372 scopus 로고
    • Polyoma small and middle T antigens and SV40 small t antigen form stable complexes with protein phosphatase 2A
    • Pallas DC, Shahrik LK, Martin BL, Jaspers S, Miller TB, Brautigan DL et al. Polyoma small and middle T antigens and SV40 small t antigen form stable complexes with protein phosphatase 2A. Cell 1990; 60: 167-176.
    • (1990) Cell , vol.60 , pp. 167-176
    • Pallas, D.C.1    Shahrik, L.K.2    Martin, B.L.3    Jaspers, S.4    Miller, T.B.5    Brautigan, D.L.6
  • 6
    • 0031784937 scopus 로고    scopus 로고
    • Signaling from polyomavirus middle T and small T defines different roles for protein phosphatase 2A
    • Mullane KP, Ratnofsky M, Cullere X, Schaffhausen B. Signaling from polyomavirus middle T and small T defines different roles for protein phosphatase 2A. Mol Cell Biol 1998; 18: 7556-7564.
    • (1998) Mol Cell Biol , vol.18 , pp. 7556-7564
    • Mullane, K.P.1    Ratnofsky, M.2    Cullere, X.3    Schaffhausen, B.4
  • 7
    • 84857133852 scopus 로고    scopus 로고
    • The role and therapeutic potential of Ser/Thr phosphatase PP2A in apoptotic signalling networks in human cancer cells
    • Janssens V, Rebollo A. The role and therapeutic potential of Ser/Thr phosphatase PP2A in apoptotic signalling networks in human cancer cells. Curr Mol Med 2012; 12: 268-287.
    • (2012) Curr Mol Med , vol.12 , pp. 268-287
    • Janssens, V.1    Rebollo, A.2
  • 8
    • 0035252894 scopus 로고    scopus 로고
    • Protein phosphatase 2A: A highly regulated family of serine/threonine phosphatases implicated in cell growth and signalling
    • Janssens V, Goris J. Protein phosphatase 2A: a highly regulated family of serine/threonine phosphatases implicated in cell growth and signalling. Biochem J 2001; 353: 417-439.
    • (2001) Biochem J , vol.353 , pp. 417-439
    • Janssens, V.1    Goris, J.2
  • 10
    • 0035754324 scopus 로고    scopus 로고
    • Protein phosphatase 2A: Variety of forms and diversity of functions
    • Lechward K, Awotunde OS, Swiatek W, Muszynska G. Protein phosphatase 2A: variety of forms and diversity of functions. Acta Biochim Pol 2001; 48: 921-933.
    • (2001) Acta Biochim Pol , vol.48 , pp. 921-933
    • Lechward, K.1    Awotunde, O.S.2    Swiatek, W.3    Muszynska, G.4
  • 11
    • 84878431342 scopus 로고    scopus 로고
    • Phosphatase: PP2A structural importance, regulation and its aberrant expression in cancer
    • Seshacharyulu P, Pandey P, Datta K, Batra SK. Phosphatase: PP2A structural importance, regulation and its aberrant expression in cancer. Cancer Lett 2013; 335: 9-18.
    • (2013) Cancer Lett , vol.335 , pp. 9-18
    • Seshacharyulu, P.1    Pandey, P.2    Datta, K.3    Batra, S.K.4
  • 12
    • 84872761033 scopus 로고    scopus 로고
    • The biogenesis of active protein phosphatase 2A holoenzymes: A tightly regulated process creating phosphatase specificity
    • Sents W, Ivanova E, Lambrecht C, Haesen D, Janssens V. The biogenesis of active protein phosphatase 2A holoenzymes: a tightly regulated process creating phosphatase specificity. FEBS J 2013; 280: 644-661.
    • (2013) FEBS J , vol.280 , pp. 644-661
    • Sents, W.1    Ivanova, E.2    Lambrecht, C.3    Haesen, D.4    Janssens, V.5
  • 13
    • 0035840446 scopus 로고    scopus 로고
    • Role of serine/threonine protein phosphatase 2A in cancer
    • Schonthal AH. Role of serine/threonine protein phosphatase 2A in cancer. Cancer Lett 2001; 170: 1-13.
    • (2001) Cancer Lett , vol.170 , pp. 1-13
    • Schonthal, A.H.1
  • 14
    • 0037424266 scopus 로고    scopus 로고
    • The formation and activity of PP2A holoenzymes do not depend on the isoform of the catalytic subunit
    • Zhou J, Pham HT, Walter G. The formation and activity of PP2A holoenzymes do not depend on the isoform of the catalytic subunit. J Biol Chem 2003; 278: 8617-8622.
    • (2003) J Biol Chem , vol.278 , pp. 8617-8622
    • Zhou, J.1    Pham, H.T.2    Walter, G.3
  • 15
    • 80054978697 scopus 로고    scopus 로고
    • Comparisons between murine polyomavirus and Simian virus 40 show significant differences in small T antigen function
    • Andrabi S, Hwang JH, Choe JK, Roberts TM. Schaffhausen BS. Comparisons between murine polyomavirus and Simian virus 40 show significant differences in small T antigen function. J Virol 2011; 85: 10649-10658.
    • (2011) J Virol , vol.85 , pp. 10649-10658
    • Andrabi, S.1    Hwang, J.H.2    Choe, J.K.3    Roberts, T.M.4    Schaffhausen, B.S.5
  • 16
    • 37649024728 scopus 로고    scopus 로고
    • Protein phosphatase 2A regulates life and death decisions via Akt in a context-dependent manner
    • Andrabi S, Gjoerup OV, Kean JA, Roberts TM. Schaffhausen B. Protein phosphatase 2A regulates life and death decisions via Akt in a context-dependent manner. Proc Natl Acad Sci USA 2007; 104: 19011-19016.
    • (2007) Proc Natl Acad Sci USA , vol.104 , pp. 19011-19016
    • Andrabi, S.1    Gjoerup, O.V.2    Kean, J.A.3    Roberts, T.M.4    Schaffhausen, B.5
  • 17
    • 0021242359 scopus 로고
    • Construction and functional characterization of polyomavirus genomes that separately encode the three early proteins
    • Zhu ZY, Veldman GM, Cowie A, Carr A, Schaffhausen B, Kamen R. Construction and functional characterization of polyomavirus genomes that separately encode the three early proteins. J Virol 1984; 51: 170-180.
    • (1984) J Virol , vol.51 , pp. 170-180
    • Zhu, Z.Y.1    Veldman, G.M.2    Cowie, A.3    Carr, A.4    Schaffhausen, B.5    Kamen, R.6
  • 18
    • 0025914438 scopus 로고
    • Mitotic arrest and enhanced nuclear protein phosphorylation in human leukemia K562 cells by okadaic acid, a potent protein phosphatase inhibitor and tumor promoter
    • Zheng B, Woo CF, Kuo JF. Mitotic arrest and enhanced nuclear protein phosphorylation in human leukemia K562 cells by okadaic acid, a potent protein phosphatase inhibitor and tumor promoter. J Biol Chem 1991; 266: 10031-10034.
    • (1991) J Biol Chem , vol.266 , pp. 10031-10034
    • Zheng, B.1    Woo, C.F.2    Kuo, J.F.3
  • 19
    • 34848840325 scopus 로고    scopus 로고
    • Spindle orientation in animal cell mitosis: Roles of integrin in the control of spindle axis
    • Toyoshima F, Nishida E. Spindle orientation in animal cell mitosis: roles of integrin in the control of spindle axis. J Cell Physiol 2007; 213: 407-411.
    • (2007) J Cell Physiol , vol.213 , pp. 407-411
    • Toyoshima, F.1    Nishida, E.2
  • 20
    • 0035844878 scopus 로고    scopus 로고
    • Spindle checkpoint protein Bub1 is required for kinetochore localization of Mad1, Mad2, Bub3, and CENP-E, independently of its kinase activity
    • Sharp-Baker H, Chen RH. Spindle checkpoint protein Bub1 is required for kinetochore localization of Mad1, Mad2, Bub3, and CENP-E, independently of its kinase activity. J Cell Biol 2001; 153: 1239-1250.
    • (2001) J Cell Biol , vol.153 , pp. 1239-1250
    • Sharp-Baker, H.1    Chen, R.H.2
  • 21
    • 0035694452 scopus 로고    scopus 로고
    • Kinetochore localisation and phosphorylation of the mitotic checkpoint components Bub1 and BubR1 are differentially regulated by spindle events in human cells
    • Taylor SS, Hussein D, Wang Y, Elderkin S, Morrow CJ. Kinetochore localisation and phosphorylation of the mitotic checkpoint components Bub1 and BubR1 are differentially regulated by spindle events in human cells. J Cell Sci 2001; 114: 4385-4395.
    • (2001) J Cell Sci , vol.114 , pp. 4385-4395
    • Taylor, S.S.1    Hussein, D.2    Wang, Y.3    Elderkin, S.4    Morrow, C.J.5
  • 23
    • 38849199681 scopus 로고    scopus 로고
    • Visualizing spatiotemporal dynamics of multicellular cell-cycle progression
    • Sakaue-Sawano A, Kurokawa H, Morimura T, Hanyu A, Hama H, Osawa H et al. Visualizing spatiotemporal dynamics of multicellular cell-cycle progression. Cell 2008; 132: 487-498.
    • (2008) Cell , vol.132 , pp. 487-498
    • Sakaue-Sawano, A.1    Kurokawa, H.2    Morimura, T.3    Hanyu, A.4    Hama, H.5    Osawa, H.6
  • 24
    • 34547159757 scopus 로고    scopus 로고
    • P53 mediates senescence-like arrest induced by chronic replicational stress
    • Marusyk A, Wheeler LJ, Mathews CK, DeGregori J. p53 mediates senescence-like arrest induced by chronic replicational stress. Mol Cell Biol 2007; 27: 5336-5351.
    • (2007) Mol Cell Biol , vol.27 , pp. 5336-5351
    • Marusyk, A.1    Wheeler, L.J.2    Mathews, C.K.3    Degregori, J.4
  • 25
    • 0012177733 scopus 로고    scopus 로고
    • Cyclotherapy: Protection of normal cells and unshielding of cancer cells
    • Blagosklonny MV, Darzynkiewicz Z. Cyclotherapy: protection of normal cells and unshielding of cancer cells. Cell Cycle 2002; 1: 375-382.
    • (2002) Cell Cycle , vol.1 , pp. 375-382
    • Blagosklonny, M.V.1    Darzynkiewicz, Z.2
  • 26
    • 59449098593 scopus 로고    scopus 로고
    • Lessons from polyoma middle T antigen on signaling and transformation: A DNA tumor virus contribution to the war on cancer
    • Schaffhausen BS, Roberts TM. Lessons from polyoma middle T antigen on signaling and transformation: A DNA tumor virus contribution to the war on cancer. Virology 2009; 384: 304-316.
    • (2009) Virology , vol.384 , pp. 304-316
    • Schaffhausen, B.S.1    Roberts, T.M.2
  • 29
    • 33646098667 scopus 로고    scopus 로고
    • PP2A is required for centromeric localization of Sgo1 and proper chromosome segregation
    • Tang Z, Shu H, Qi W, Mahmood NA, Mumby MC, Yu H. PP2A is required for centromeric localization of Sgo1 and proper chromosome segregation. Dev Cell 2006; 10: 575-585.
    • (2006) Dev Cell , vol.10 , pp. 575-585
    • Tang, Z.1    Shu, H.2    Qi, W.3    Mahmood, N.A.4    Mumby, M.C.5    Yu, H.6
  • 30
    • 33646819227 scopus 로고    scopus 로고
    • Protein phosphatase 2A protects centromeric sister chromatid cohesion during meiosis i
    • Riedel CG, Katis VL, Katou Y, Mori S, Itoh T, Helmhart W et al. Protein phosphatase 2A protects centromeric sister chromatid cohesion during meiosis I. Nature 2006; 441: 53-61.
    • (2006) Nature , vol.441 , pp. 53-61
    • Riedel, C.G.1    Katis, V.L.2    Katou, Y.3    Mori, S.4    Itoh, T.5    Helmhart, W.6
  • 31
    • 70849120184 scopus 로고    scopus 로고
    • Studies of haspin-depleted cells reveal that spindle-pole integrity in mitosis requires chromosome cohesion
    • Dai J, Kateneva AV, Higgins JM. Studies of haspin-depleted cells reveal that spindle-pole integrity in mitosis requires chromosome cohesion. J Cell Sci 2009; 122: 4168-4176.
    • (2009) J Cell Sci , vol.122 , pp. 4168-4176
    • Dai, J.1    Kateneva, A.V.2    Higgins, J.M.3
  • 32
    • 80053573427 scopus 로고    scopus 로고
    • Formation of stable attachments between kinetochores and microtubules depends on the B56-PP2A phosphatase
    • Foley EA, Maldonado M, Kapoor TM. Formation of stable attachments between kinetochores and microtubules depends on the B56-PP2A phosphatase. Nat Cell Biol 2011; 13: 1265-1271.
    • (2011) Nat Cell Biol , vol.13 , pp. 1265-1271
    • Foley, E.A.1    Maldonado, M.2    Kapoor, T.M.3
  • 33
    • 84876330632 scopus 로고    scopus 로고
    • MISP is a novel Plk1 substrate required for proper spindle orientation and mitotic progression
    • Zhu M, Settele F, Kotak S, Sanchez-Pulido L, Ehret L, Ponting CP et al. MISP is a novel Plk1 substrate required for proper spindle orientation and mitotic progression. J Cell Biol 2013; 200: 773-787.
    • (2013) J Cell Biol , vol.200 , pp. 773-787
    • Zhu, M.1    Settele, F.2    Kotak, S.3    Sanchez-Pulido, L.4    Ehret, L.5    Ponting, C.P.6
  • 34
    • 13444256277 scopus 로고    scopus 로고
    • Simian virus 40 small t antigen mediates conformation-dependent transfer of protein phosphatase 2A onto the androgen receptor
    • Yang CS, Vitto MJ, Busby SA, Garcia BA, Kesler CT, Gioeli D et al. Simian virus 40 small t antigen mediates conformation-dependent transfer of protein phosphatase 2A onto the androgen receptor. Mol Cell Biol 2005; 25: 1298-1308.
    • (2005) Mol Cell Biol , vol.25 , pp. 1298-1308
    • Yang, C.S.1    Vitto, M.J.2    Busby, S.A.3    Garcia, B.A.4    Kesler, C.T.5    Gioeli, D.6
  • 36
    • 0026016248 scopus 로고
    • Dephosphorylation of simian virus 40 large-T antigen and p53 protein by protein phosphatase 2A: Inhibition by small-t antigen
    • Scheidtmann KH, Mumby MC, Rundell K, Walter G. Dephosphorylation of simian virus 40 large-T antigen and p53 protein by protein phosphatase 2A: inhibition by small-t antigen. Mol Cell Biol 1991; 11: 1996-2003.
    • (1991) Mol Cell Biol , vol.11 , pp. 1996-2003
    • Scheidtmann, K.H.1    Mumby, M.C.2    Rundell, K.3    Walter, G.4
  • 37
    • 4444344407 scopus 로고    scopus 로고
    • G2 checkpoint abrogators as anticancer drugs
    • Kawabe T. G2 checkpoint abrogators as anticancer drugs. Mol Cancer Ther 2004; 3: 513-519.
    • (2004) Mol Cancer Ther , vol.3 , pp. 513-519
    • Kawabe, T.1
  • 38
    • 34648832173 scopus 로고    scopus 로고
    • FTY720, a new alternative for treating blast crisis chronic myelogenous leukemia and Philadelphia chromosome-positive acute lymphocytic leukemia
    • Neviani P, Santhanam R, Oaks JJ, Eiring AM, Notari M, Blaser BW et al. FTY720, a new alternative for treating blast crisis chronic myelogenous leukemia and Philadelphia chromosome-positive acute lymphocytic leukemia. J Clin Invest 2007; 117: 2408-2421.
    • (2007) J Clin Invest , vol.117 , pp. 2408-2421
    • Neviani, P.1    Santhanam, R.2    Oaks, J.J.3    Eiring, A.M.4    Notari, M.5    Blaser, B.W.6
  • 39
    • 84887057719 scopus 로고    scopus 로고
    • Protein phosphatase 2A isoforms utilizing Abeta scaffolds regulate differentiation through control of Akt protein
    • Hwang JH, Jiang T, Kulkarni S, Faure N. Schaffhausen BS. Protein phosphatase 2A isoforms utilizing Abeta scaffolds regulate differentiation through control of Akt protein. J Biol Chem 2013; 288: 32064-32073.
    • (2013) J Biol Chem , vol.288 , pp. 32064-32073
    • Hwang, J.H.1    Jiang, T.2    Kulkarni, S.3    Faure, N.4    Schaffhausen, B.S.5
  • 40
    • 0036032778 scopus 로고    scopus 로고
    • Regulators of serine/threonine protein phosphatases at the dawn of a clinical era?
    • Honkanen RE, Golden T. Regulators of serine/threonine protein phosphatases at the dawn of a clinical era? Curr Med Chem 2002; 9: 2055-2075.
    • (2002) Curr Med Chem , vol.9 , pp. 2055-2075
    • Honkanen, R.E.1    Golden, T.2
  • 41
    • 34447275978 scopus 로고    scopus 로고
    • Small-molecule inhibitors of ser/thr protein phosphatases: Specificity, use and common forms of abuse
    • Swingle M, Ni L, Honkanen RE. Small-molecule inhibitors of ser/thr protein phosphatases: specificity, use and common forms of abuse. Methods Mol Biol 2007; 365: 23-38.
    • (2007) Methods Mol Biol , vol.365 , pp. 23-38
    • Swingle, M.1    Ni, L.2    Honkanen, R.E.3
  • 42
    • 0034807394 scopus 로고    scopus 로고
    • Overexpression of simian virus 40 small-T antigen blocks centrosome function and mitotic progression in human fibroblasts
    • Gaillard S, Fahrbach KM, Parkati R, Rundell K. Overexpression of simian virus 40 small-T antigen blocks centrosome function and mitotic progression in human fibroblasts. J Virol 2001; 75: 9799-9807.
    • (2001) J Virol , vol.75 , pp. 9799-9807
    • Gaillard, S.1    Fahrbach, K.M.2    Parkati, R.3    Rundell, K.4
  • 43
    • 0034847978 scopus 로고    scopus 로고
    • Induction of p53-independent apoptosis by simian virus 40 small t antigen
    • Gjoerup O, Zaveri D, Roberts TM. Induction of p53-independent apoptosis by simian virus 40 small t antigen. J Virol 2001; 75: 9142-9155.
    • (2001) J Virol , vol.75 , pp. 9142-9155
    • Gjoerup, O.1    Zaveri, D.2    Roberts, T.M.3
  • 44
    • 69249229734 scopus 로고    scopus 로고
    • The adenovirus E4orf4 protein induces G2/M arrest and cell death by blocking protein phosphatase 2A activity regulated by the B55 subunit
    • Li S, Brignole C, Marcellus R, Thirlwell S, Binda O, McQuoid MJ et al. The adenovirus E4orf4 protein induces G2/M arrest and cell death by blocking protein phosphatase 2A activity regulated by the B55 subunit. J Virol 2009; 83: 8340-8352.
    • (2009) J Virol , vol.83 , pp. 8340-8352
    • Li, S.1    Brignole, C.2    Marcellus, R.3    Thirlwell, S.4    Binda, O.5    McQuoid, M.J.6
  • 45
    • 59649087879 scopus 로고    scopus 로고
    • The adenovirus E4orf4 protein induces growth arrest and mitotic catastrophe in H1299 human lung carcinoma cells
    • Li S, Szymborski A, Miron MJ, Marcellus R, Binda O, Lavoie JN et al. The adenovirus E4orf4 protein induces growth arrest and mitotic catastrophe in H1299 human lung carcinoma cells. Oncogene 2009; 28: 390-400.
    • (2009) Oncogene , vol.28 , pp. 390-400
    • Li, S.1    Szymborski, A.2    Miron, M.J.3    Marcellus, R.4    Binda, O.5    Lavoie, J.N.6
  • 47
    • 25444440871 scopus 로고    scopus 로고
    • Modulation of the ARF-p53 pathway by the small DNA tumor viruses
    • O'Shea CC, Fried M. Modulation of the ARF-p53 pathway by the small DNA tumor viruses. Cell Cycle 2005; 4: 449-452.
    • (2005) Cell Cycle , vol.4 , pp. 449-452
    • O'Shea, C.C.1    Fried, M.2
  • 48
    • 33846584645 scopus 로고    scopus 로고
    • Mitotic arrest and cell fate: Why and how mitotic inhibition of transcription drives mutually exclusive events
    • Blagosklonny MV. Mitotic arrest and cell fate: why and how mitotic inhibition of transcription drives mutually exclusive events. Cell Cycle 2007; 6: 70-74.
    • (2007) Cell Cycle , vol.6 , pp. 70-74
    • Blagosklonny, M.V.1
  • 50
    • 84879804348 scopus 로고    scopus 로고
    • Nourseothricin N-acetyl transferase: A positive selection marker for mammalian cells
    • Kochupurakkal BS, Iglehart JD. Nourseothricin N-acetyl transferase: a positive selection marker for mammalian cells. PLoS ONE 2013; 8: e68509.
    • (2013) PLoS ONE , vol.8 , pp. e68509
    • Kochupurakkal, B.S.1    Iglehart, J.D.2
  • 51
    • 33750428233 scopus 로고    scopus 로고
    • Regulation of mitotic chromosome cohesion by Haspin and Aurora B
    • Dai J, Sullivan BA, Higgins JM. Regulation of mitotic chromosome cohesion by Haspin and Aurora B. Dev Cell 2006; 11: 741-750.
    • (2006) Dev Cell , vol.11 , pp. 741-750
    • Dai, J.1    Sullivan, B.A.2    Higgins, J.M.3
  • 52
    • 0038485575 scopus 로고    scopus 로고
    • TGF-beta1 suppresses apoptosis via differential regulation of MAP kinases and ceramide production
    • Chen HH, Zhao S, Song JG. TGF-beta1 suppresses apoptosis via differential regulation of MAP kinases and ceramide production. Cell Death Differ 2003; 10: 516-527
    • (2003) Cell Death Differ , vol.10 , pp. 516-527
    • Chen, H.H.1    Zhao, S.2    Song, J.G.3


* 이 정보는 Elsevier사의 SCOPUS DB에서 KISTI가 분석하여 추출한 것입니다.