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Volumn 82, Issue 10, 2008, Pages 4898-4907

Efficient and specific rescue of human immunodeficiency virus type 1 budding defects by a Nedd4-like ubiquitin ligase

Author keywords

[No Author keywords available]

Indexed keywords

GAG PROTEIN; ISOPROTEIN; MUTANT PROTEIN; PROTEIN ALIX; THIOESTER; TUMOR SUSCEPTIBILITY GENE 101 PROTEIN; UBIQUITIN PROTEIN LIGASE NEDD4; VIRUS PROTEIN; NEDD4 UBIQUITIN PROTEIN LIGASES; P6 GAG PROTEIN, HUMAN IMMUNODEFICIENCY VIRUS 1; UBIQUITIN; UBIQUITIN PROTEIN LIGASE;

EID: 43249110449     PISSN: 0022538X     EISSN: None     Source Type: Journal    
DOI: 10.1128/JVI.02675-07     Document Type: Article
Times cited : (93)

References (56)
  • 1
    • 0034088332 scopus 로고    scopus 로고
    • Efficient particle production by minimal Gag constructs which retain the carboxy-terminal domain of human immunodeficiency virus type 1 capsid-p2 and a late assembly domain
    • Accola, M. A., B. Strack, and H. G. Gottlinger. 2000. Efficient particle production by minimal Gag constructs which retain the carboxy-terminal domain of human immunodeficiency virus type 1 capsid-p2 and a late assembly domain. J. Virol. 74:5395-5402.
    • (2000) J. Virol , vol.74 , pp. 5395-5402
    • Accola, M.A.1    Strack, B.2    Gottlinger, H.G.3
  • 2
    • 12444347534 scopus 로고    scopus 로고
    • A protein's final ESCRT
    • Babst, M. 2005. A protein's final ESCRT. Traffic 6:2-9.
    • (2005) Traffic , vol.6 , pp. 2-9
    • Babst, M.1
  • 3
    • 0036696804 scopus 로고    scopus 로고
    • Escrt-III: An endosome-associatcd heterooligomeric protein complex required for mvb sorting
    • Babst, M., D. J. Katzmann, E. J. Estepa-Sabal, T. Meerloo, and S. D. Emr. 2002. Escrt-III: an endosome-associatcd heterooligomeric protein complex required for mvb sorting. Dev. Cell 3:271-282.
    • (2002) Dev. Cell , vol.3 , pp. 271-282
    • Babst, M.1    Katzmann, D.J.2    Estepa-Sabal, E.J.3    Meerloo, T.4    Emr, S.D.5
  • 4
    • 29144474443 scopus 로고    scopus 로고
    • Late budding domains and host proteins in enveloped virus release
    • Bieniasz, P. D. 2006. Late budding domains and host proteins in enveloped virus release. Virology 344:55-63.
    • (2006) Virology , vol.344 , pp. 55-63
    • Bieniasz, P.D.1
  • 5
    • 0031680643 scopus 로고    scopus 로고
    • The C-terminal half of the human immunodeficiency virus type 1 Gag precursor is sufficient for efficient particle assembly
    • Borsetti, A., A. Ohagen, and H. G. Gottlinger. 1998. The C-terminal half of the human immunodeficiency virus type 1 Gag precursor is sufficient for efficient particle assembly. J. Virol. 72:9313-9317.
    • (1998) J. Virol , vol.72 , pp. 9313-9317
    • Borsetti, A.1    Ohagen, A.2    Gottlinger, H.G.3
  • 6
    • 0242331750 scopus 로고    scopus 로고
    • Bouamr, F., J. A. Melillo, M. Q. Wang, K. Nagashima, M. de Los Santos, A. Rein, and S. P. Goff. 2003. PPPYEPTAP motif is the late domain of human T-cell leukemia virus type 1 Gag and mediates its functional interaction with cellular proteins Nedd4 and Tsg101. J. Virol. 77:11882-11895. [Author's correction. 78:4383, 2004.]
    • Bouamr, F., J. A. Melillo, M. Q. Wang, K. Nagashima, M. de Los Santos, A. Rein, and S. P. Goff. 2003. PPPYEPTAP motif is the late domain of human T-cell leukemia virus type 1 Gag and mediates its functional interaction with cellular proteins Nedd4 and Tsg101. J. Virol. 77:11882-11895. [Author's correction. 78:4383, 2004.]
  • 7
    • 0030858496 scopus 로고    scopus 로고
    • Characterization of the WW domain of human yes-associated protein and its polvproline-eontaining ligands
    • Chen, H. I., A. Einbond, S. J. Kwak, H. Linn, E. Koepf, S. Peterson, J. W. Kelly, and M. Sudol. 1997, Characterization of the WW domain of human yes-associated protein and its polvproline-eontaining ligands. J. Biol. Chem. 272:17070-17077.
    • (1997) J. Biol. Chem , vol.272 , pp. 17070-17077
    • Chen, H.I.1    Einbond, A.2    Kwak, S.J.3    Linn, H.4    Koepf, E.5    Peterson, S.6    Kelly, J.W.7    Sudol, M.8
  • 8
    • 0026672676 scopus 로고
    • Macrophage-tropic human immunodeficiency virus isolates from different patients exhibit unusual V3 envelope sequence homogeneity in comparison with T-Cell-tropic isolates: Definition of critical amino acids involved in cell tropism
    • Chesebro, B., K. Wehrly, J. Nishio, and S. Perryman, 1992. Macrophage-tropic human immunodeficiency virus isolates from different patients exhibit unusual V3 envelope sequence homogeneity in comparison with T-Cell-tropic isolates: definition of critical amino acids involved in cell tropism. J. Virol. 66:6547-6554.
    • (1992) J. Virol , vol.66 , pp. 6547-6554
    • Chesebro, B.1    Wehrly, K.2    Nishio, J.3    Perryman, S.4
  • 9
    • 43249121457 scopus 로고    scopus 로고
    • NEDD4L overexpression rescues the release and infectivity of human immunodeficiency virus type 1 constructs lacking PTAP and YPXL late domains
    • Chung, H.-Y., E. Morita, U. von Schwedler, B. Müller, H.-G. Kräusslich, and W. I. Sundquist. 2008. NEDD4L overexpression rescues the release and infectivity of human immunodeficiency virus type 1 constructs lacking PTAP and YPXL late domains. J. Virol. 82:4884-4897.
    • (2008) J. Virol , vol.82 , pp. 4884-4897
    • Chung, H.-Y.1    Morita, E.2    von Schwedler, U.3    Müller, B.4    Kräusslich, H.-G.5    Sundquist, W.I.6
  • 10
    • 9744221135 scopus 로고    scopus 로고
    • Retrovirus budding
    • Demirov, D. G., and E. O. Freed. 2004. Retrovirus budding. Virus Res. 106:87-102.
    • (2004) Virus Res , vol.106 , pp. 87-102
    • Demirov, D.G.1    Freed, E.O.2
  • 11
    • 0027997831 scopus 로고
    • Functional domains of the capsid protein of human immunodeficiency virus type 1
    • Dorfman, T., A. Bukovsky, A. Ohagen, S. Hoglund, and H. G. Gottlinger. 1994. Functional domains of the capsid protein of human immunodeficiency virus type 1. J. Virol. 68:8180-8187.
    • (1994) J. Virol , vol.68 , pp. 8180-8187
    • Dorfman, T.1    Bukovsky, A.2    Ohagen, A.3    Hoglund, S.4    Gottlinger, H.G.5
  • 13
    • 33847355934 scopus 로고    scopus 로고
    • Structural and biochemical studies of ALIX/AIP1 and its role in retrovirus budding
    • Fisher, R. D., H. Y. Chung, Q. Zhai, H. Robinson, W. I. Sundquist, and C. P. Hill. 2007. Structural and biochemical studies of ALIX/AIP1 and its role in retrovirus budding. Cell 128:841-852.
    • (2007) Cell , vol.128 , pp. 841-852
    • Fisher, R.D.1    Chung, H.Y.2    Zhai, Q.3    Robinson, H.4    Sundquist, W.I.5    Hill, C.P.6
  • 15
    • 0026317877 scopus 로고
    • Effect of mutations affecting the p6 gag protein on human immunodeficiency virus particle release
    • Gottlinger, H. G., T. Dorfman, J. G. Sodroski, and W. A. Haseltine. 1991. Effect of mutations affecting the p6 gag protein on human immunodeficiency virus particle release. Proc. Natl. Acad. Sci. USA 88:3195-3199.
    • (1991) Proc. Natl. Acad. Sci. USA , vol.88 , pp. 3195-3199
    • Gottlinger, H.G.1    Dorfman, T.2    Sodroski, J.G.3    Haseltine, W.A.4
  • 16
    • 0042389562 scopus 로고    scopus 로고
    • The Mason-Pfizer monkey virus PPPY and PSAP motifs hoth contribute to virus release
    • Gottwein, E., J. Bodem, B. Muller, A. Schmechel, H. Zentgraf, and H. G. Krausslich. 2003. The Mason-Pfizer monkey virus PPPY and PSAP motifs hoth contribute to virus release. J. Virol. 77:9474-9485.
    • (2003) J. Virol , vol.77 , pp. 9474-9485
    • Gottwein, E.1    Bodem, J.2    Muller, B.3    Schmechel, A.4    Zentgraf, H.5    Krausslich, H.G.6
  • 17
    • 33745234847 scopus 로고    scopus 로고
    • Cumulative mutations of ubiquitin acceptor sites in human immunodeficiency virus type 1 Gag cause a late budding defect
    • Gottwein, E., S. Jager, A. Habermann, and H. G. Krausslich. 2006. Cumulative mutations of ubiquitin acceptor sites in human immunodeficiency virus type 1 Gag cause a late budding defect. J. Virol. 80:6267-6275.
    • (2006) J. Virol , vol.80 , pp. 6267-6275
    • Gottwein, E.1    Jager, S.2    Habermann, A.3    Krausslich, H.G.4
  • 18
    • 0034610295 scopus 로고    scopus 로고
    • A PPxY motif within the VP40 protein of Ebola virus interacts physically and functionally with a ubiquitin ligase: Implications for filovirus budding
    • Harty, R. N., M. E. Brown, G. Wang, J. Huibregtse, and F. P. Hayes. 2000. A PPxY motif within the VP40 protein of Ebola virus interacts physically and functionally with a ubiquitin ligase: implications for filovirus budding. Proc. Natl. Acad. Sci. USA 97:13871-13876.
    • (2000) Proc. Natl. Acad. Sci. USA , vol.97 , pp. 13871-13876
    • Harty, R.N.1    Brown, M.E.2    Wang, G.3    Huibregtse, J.4    Hayes, F.P.5
  • 21
    • 0028971135 scopus 로고
    • p6Gag is required for particle production from full-length human immunodeficiency virus type 1 molecular clones expressing protease
    • Huang, M., J. M. Orenstein, M. A. Martin, and E. O. Freed. 1995. p6Gag is required for particle production from full-length human immunodeficiency virus type 1 molecular clones expressing protease. J. Virol. 69:6810-6818.
    • (1995) J. Virol , vol.69 , pp. 6810-6818
    • Huang, M.1    Orenstein, J.M.2    Martin, M.A.3    Freed, E.O.4
  • 22
    • 33646010475 scopus 로고    scopus 로고
    • The ESCRT complexes: Structure and mechanism of a membrane-trafficking network
    • Hurley, J. H., and S. D. Emr. 2006. The ESCRT complexes: structure and mechanism of a membrane-trafficking network. Annu. Rev. Biophys. Biomol. Struct. 35:277-298.
    • (2006) Annu. Rev. Biophys. Biomol. Struct , vol.35 , pp. 277-298
    • Hurley, J.H.1    Emr, S.D.2
  • 23
    • 1842591240 scopus 로고    scopus 로고
    • The Nedd4 family of E3 ubiquitin ligases: Functional diversity within a common modular architecture
    • Ingham, R. J., G. Gish, and T. Pawson. 2004. The Nedd4 family of E3 ubiquitin ligases: functional diversity within a common modular architecture. Oncogene 23:1972-1984.
    • (2004) Oncogene , vol.23 , pp. 1972-1984
    • Ingham, R.J.1    Gish, G.2    Pawson, T.3
  • 24
    • 0142025380 scopus 로고    scopus 로고
    • Alternate promoters and variable splicing lead to hNedd4-2 isoforms with a C2 domain and varying number of WW domains
    • Itani, O. A., J. R. Campbell, J. Herrero, P. M. Snyder, and C. P. Thomas. 2003. Alternate promoters and variable splicing lead to hNedd4-2 isoforms with a C2 domain and varying number of WW domains. Am. J. Physiol. Renal Physiol. 285:F916-F929.
    • (2003) Am. J. Physiol. Renal Physiol , vol.285
    • Itani, O.A.1    Campbell, J.R.2    Herrero, J.3    Snyder, P.M.4    Thomas, C.P.5
  • 25
    • 0034819622 scopus 로고    scopus 로고
    • Distinct characteristics of two human Nedd4 proteins with respect to epithelial Na(+) channel regulation
    • Kamynina, E., C. Tauxe, and O. Staub. 2001. Distinct characteristics of two human Nedd4 proteins with respect to epithelial Na(+) channel regulation. Am. J. Physiol. Renal Physiol. 281:F469-F477.
    • (2001) Am. J. Physiol. Renal Physiol , vol.281
    • Kamynina, E.1    Tauxe, C.2    Staub, O.3
  • 26
    • 18544383164 scopus 로고    scopus 로고
    • Ligand-independent degradation of epidermal growth factor receptor involves receptor ubiquitylation and Hgs, an adaptor whose ubiquitin-interacting motif targets ubiquitylation by Nedd4
    • Katz, M., K. Shtiegman, P. Tal-Or, L. Yakir, Y. Mosesson, D. Harari, Y. Machluf, H. Asao, T. Jovin, K. Sugamura, and Y. Yarden. 2002. Ligand-independent degradation of epidermal growth factor receptor involves receptor ubiquitylation and Hgs, an adaptor whose ubiquitin-interacting motif targets ubiquitylation by Nedd4. Traffic 3:740-751.
    • (2002) Traffic , vol.3 , pp. 740-751
    • Katz, M.1    Shtiegman, K.2    Tal-Or, P.3    Yakir, L.4    Mosesson, Y.5    Harari, D.6    Machluf, Y.7    Asao, H.8    Jovin, T.9    Sugamura, K.10    Yarden, Y.11
  • 27
    • 0034517389 scopus 로고    scopus 로고
    • Smad7 binds to Smurf2 to form an E3 ubiquitin ligase that targets the TGF beta receptor for degradation
    • Kavsak, P., R. K. Rasmussen, C. G. Causing, S. Bonni, H. Zhu, G. H. Thomsen, and J. L. Wrana. 2000. Smad7 binds to Smurf2 to form an E3 ubiquitin ligase that targets the TGF beta receptor for degradation. Mol. Cell 6:1365-1375.
    • (2000) Mol. Cell , vol.6 , pp. 1365-1375
    • Kavsak, P.1    Rasmussen, R.K.2    Causing, C.G.3    Bonni, S.4    Zhu, H.5    Thomsen, G.H.6    Wrana, J.L.7
  • 28
    • 0035949489 scopus 로고    scopus 로고
    • Proteins related to the Nedd4 family of ubiquitin protein ligases interact with the L domain of Rous sarcoma virus and are required for gag budding from cells
    • Kikonyogo, A., F. Bouamr, M. L. Vana, Y. Xiang, A. Aiyar, C. Carter, and J. Leis. 2001. Proteins related to the Nedd4 family of ubiquitin protein ligases interact with the L domain of Rous sarcoma virus and are required for gag budding from cells. Proc. Natl. Acad. Sci. USA 98:11199-11204.
    • (2001) Proc. Natl. Acad. Sci. USA , vol.98 , pp. 11199-11204
    • Kikonyogo, A.1    Bouamr, F.2    Vana, M.L.3    Xiang, Y.4    Aiyar, A.5    Carter, C.6    Leis, J.7
  • 29
    • 0040017910 scopus 로고    scopus 로고
    • Function of WW domains as phosphoserine- or phosphothreonine-binding modules
    • Lu, P. J., X. Z. Zhou, M. Shen, and K. P. Lu. 1999. Function of WW domains as phosphoserine- or phosphothreonine-binding modules. Science 283:1325-1328.
    • (1999) Science , vol.283 , pp. 1325-1328
    • Lu, P.J.1    Zhou, X.Z.2    Shen, M.3    Lu, K.P.4
  • 30
    • 34548580258 scopus 로고    scopus 로고
    • The role of ubiquitin in retroviral egress
    • Martin-Serrano, J. 2007. The role of ubiquitin in retroviral egress. Traffic 8:1297-1303.
    • (2007) Traffic , vol.8 , pp. 1297-1303
    • Martin-Serrano, J.1
  • 31
    • 12144258074 scopus 로고    scopus 로고
    • HECT ubiquitin ligases link viral and cellular PPXY motifs to the vacuolar protein-sorting pathway
    • Martin-Serrano, J., S. W. Eastman, W. Chung, and P. D. Bieniasz. 2005. HECT ubiquitin ligases link viral and cellular PPXY motifs to the vacuolar protein-sorting pathway. J. Cell Biol. 168:89-101.
    • (2005) J. Cell Biol , vol.168 , pp. 89-101
    • Martin-Serrano, J.1    Eastman, S.W.2    Chung, W.3    Bieniasz, P.D.4
  • 32
    • 0142123069 scopus 로고    scopus 로고
    • Divergent retroviral late-budding domains recruit vacuolar protein sorting factors by using alternative adaptor proteins
    • Martin-Serrano, J., A. Yarovoy, D. Perez-Caballero, and P. D. Bieniasz. 2003. Divergent retroviral late-budding domains recruit vacuolar protein sorting factors by using alternative adaptor proteins. Proc. Natl. Acad. Sci. USA 100:12414-12419.
    • (2003) Proc. Natl. Acad. Sci. USA , vol.100 , pp. 12414-12419
    • Martin-Serrano, J.1    Yarovoy, A.2    Perez-Caballero, D.3    Bieniasz, P.D.4
  • 34
    • 0037169521 scopus 로고    scopus 로고
    • Identification of developmentally expressed proteins that functionally interact with Nedd4 ubiquitin ligase
    • Murillas, R., K. S. Simms, S. Hatakeyama, A. M. Weissman, and M. R. Kuehn. 2002. Identification of developmentally expressed proteins that functionally interact with Nedd4 ubiquitin ligase. J. Biol. Chem. 277:2897-2907.
    • (2002) J. Biol. Chem , vol.277 , pp. 2897-2907
    • Murillas, R.1    Simms, K.S.2    Hatakeyama, S.3    Weissman, A.M.4    Kuehn, M.R.5
  • 35
    • 0034610238 scopus 로고    scopus 로고
    • Ott, D. E., L. V. Coren, E. N. Chertova, T. D. Gagliardi, and U. Schubert. 2000. Ubiquitination of HIV-1 and MuLV Gag. Virology 278:111-121.
    • Ott, D. E., L. V. Coren, E. N. Chertova, T. D. Gagliardi, and U. Schubert. 2000. Ubiquitination of HIV-1 and MuLV Gag. Virology 278:111-121.
  • 37
    • 38349174466 scopus 로고    scopus 로고
    • Human immunodeficiency virus type 1 Gag engages the Bro1 domain of ALIX/AIP1 through nucleocapsid
    • Popov, S., E. Popova, M. Inoue, and H. G. Gottlinger. 2008. Human immunodeficiency virus type 1 Gag engages the Bro1 domain of ALIX/AIP1 through nucleocapsid. J. Virol. 82:1389-1398.
    • (2008) J. Virol , vol.82 , pp. 1389-1398
    • Popov, S.1    Popova, E.2    Inoue, M.3    Gottlinger, H.G.4
  • 38
    • 0141717669 scopus 로고    scopus 로고
    • The M-PMV cytoplasmic targeting-retention signal directs nascent Gag polypeptides to a pericentriolar region of the cell
    • Sfakianos, J. N., R. A. LaCasse, and E. Hunter. 2003. The M-PMV cytoplasmic targeting-retention signal directs nascent Gag polypeptides to a pericentriolar region of the cell. Traffic 4:660-670.
    • (2003) Traffic , vol.4 , pp. 660-670
    • Sfakianos, J.N.1    LaCasse, R.A.2    Hunter, E.3
  • 39
    • 33745089231 scopus 로고    scopus 로고
    • Regulation of functional diversity within the Nedd4 family by accessory and adaptor proteins
    • Shearwin-Whyatt, L., H. E. Dalton, N. Foot, and S. Kumar. 2006. Regulation of functional diversity within the Nedd4 family by accessory and adaptor proteins. Bioessays 28:617-628.
    • (2006) Bioessays , vol.28 , pp. 617-628
    • Shearwin-Whyatt, L.1    Dalton, H.E.2    Foot, N.3    Kumar, S.4
  • 40
  • 41
    • 0035958908 scopus 로고    scopus 로고
    • Multiple WW domains, but not the C2 domain, are required for inhibition of the epithelial Na+ channel by human Nedd4
    • Snyder, P. M., D. R. Olson, F. J. McDonald, and D. B. Bucher. 2001. Multiple WW domains, but not the C2 domain, are required for inhibition of the epithelial Na+ channel by human Nedd4. J. Biol. Chem. 276:28321- 28326.
    • (2001) J. Biol. Chem , vol.276 , pp. 28321-28326
    • Snyder, P.M.1    Olson, D.R.2    McDonald, F.J.3    Bucher, D.B.4
  • 44
    • 0141844660 scopus 로고    scopus 로고
    • AIP1/ALIX is a binding partner for HIV-1 p6 and EIAV p9 functioning in virus budding
    • Strack, B., A. Calistri, S. Craig, E. Popova, and H. G. Gottlinger. 2003. AIP1/ALIX is a binding partner for HIV-1 p6 and EIAV p9 functioning in virus budding. Cell 114:689-699.
    • (2003) Cell , vol.114 , pp. 689-699
    • Strack, B.1    Calistri, A.2    Craig, S.3    Popova, E.4    Gottlinger, H.G.5
  • 45
    • 0036094492 scopus 로고    scopus 로고
    • Late assembly domain function can exhibit context dependence and involves ubiquitin residues implicated in endocytosis
    • Strack, B., A. Calistri, and H. G. Gottlinger. 2002. Late assembly domain function can exhibit context dependence and involves ubiquitin residues implicated in endocytosis. J. Virol. 76:5472-5479.
    • (2002) J. Virol , vol.76 , pp. 5472-5479
    • Strack, B.1    Calistri, A.2    Gottlinger, H.G.3
  • 48
    • 34249943479 scopus 로고    scopus 로고
    • Potent rescue of human immunodeficiency virus type 1 late domain mutants by ALIX/AIP1 depends on its CHMP4 binding site
    • Usami, Y., S. Popov, and H. G. Gottlinger. 2007. Potent rescue of human immunodeficiency virus type 1 late domain mutants by ALIX/AIP1 depends on its CHMP4 binding site. J. Virol. 81:6614-6622.
    • (2007) J. Virol , vol.81 , pp. 6614-6622
    • Usami, Y.1    Popov, S.2    Gottlinger, H.G.3
  • 53
    • 0031968983 scopus 로고    scopus 로고
    • A proline-rich motif (PPPY) in the Gag polyprotcin of Mason-Pfizer monkey virus plays a maturation-independent role in virion release
    • Yasuda, J., and E. Hunter. 1998. A proline-rich motif (PPPY) in the Gag polyprotcin of Mason-Pfizer monkey virus plays a maturation-independent role in virion release. J. Virol. 72:4095-4103.
    • (1998) J. Virol , vol.72 , pp. 4095-4103
    • Yasuda, J.1    Hunter, E.2
  • 54
    • 0036023945 scopus 로고    scopus 로고
    • Functional involvement of a novel Nedd4-like ubiquitin ligase on retrovirus budding
    • Yasuda, J., E. Hunter, M. Nakao, and H. Shida. 2002. Functional involvement of a novel Nedd4-like ubiquitin ligase on retrovirus budding. EMBO Rep. 3:636-640.
    • (2002) EMBO Rep , vol.3 , pp. 636-640
    • Yasuda, J.1    Hunter, E.2    Nakao, M.3    Shida, H.4
  • 56
    • 84881193950 scopus 로고    scopus 로고
    • Ubiquitin-dependent virus particle budding without viral protein ubiquitination
    • Zhadina, M., M. O. McClure, M. C. Johnson, and P. D. Bieniasz. 2007. Ubiquitin-dependent virus particle budding without viral protein ubiquitination. Proc. Natl. Acad. Sci. USA 104:20031-20036.
    • (2007) Proc. Natl. Acad. Sci. USA , vol.104 , pp. 20031-20036
    • Zhadina, M.1    McClure, M.O.2    Johnson, M.C.3    Bieniasz, P.D.4


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