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Volumn 432, Issue 3, 2010, Pages 461-472

Functional complexes between YAP2 and ZO-2 are PDZ domain-dependent, and regulate YAP2 nuclear localization and signalling

Author keywords

Hippo pathway; Nuclear translocation; PDZ domain; Yes kinase associated protein 2 (YAP2); Zona occludens protein

Indexed keywords

HIPPO PATHWAY; NUCLEAR TRANSLOCATIONS; PDZ DOMAINS; YES KINASE-ASSOCIATED PROTEIN 2 (YAP2); ZONA OCCLUDENS;

EID: 78649892579     PISSN: 02646021     EISSN: 14708728     Source Type: Journal    
DOI: 10.1042/BJ20100870     Document Type: Article
Times cited : (174)

References (55)
  • 1
    • 33847206401 scopus 로고    scopus 로고
    • The Salvador-Warts-Hippo pathway: An emerging tumour-suppressor network
    • Harvey, K. and Tapon, N. (2007) The Salvador-Warts-Hippo pathway: an emerging tumour-suppressor network. Nat. Rev. Cancer 7, 182-191
    • (2007) Nat. Rev. Cancer , vol.7 , pp. 182-191
    • Harvey, K.1    Tapon, N.2
  • 2
    • 0028283579 scopus 로고
    • Yes-associated protein (YAP65) is a proline-rich phosphoprotein that binds to the SH3 domain of the Yes proto-oncogene product
    • Sudol, M. (1994) Yes-associated protein (YAP65) is a proline-rich phosphoprotein that binds to the SH3 domain of the Yes proto-oncogene product. Oncogene 9, 2145-2152
    • (1994) Oncogene , vol.9 , pp. 2145-2152
    • Sudol, M.1
  • 3
    • 0035957958 scopus 로고    scopus 로고
    • Yes-associated protein and p53-binding protein-2 interact through their WW and SH3 domains
    • Espanel, X. and Sudol, M. (2001) Yes-associated protein and p53-binding protein-2 interact through their WW and SH3 domains. J. Biol. Chem. 276, 14514-14523
    • (2001) J. Biol. Chem. , vol.276 , pp. 14514-14523
    • Espanel, X.1    Sudol, M.2
  • 4
    • 0042858208 scopus 로고    scopus 로고
    • WW domain-containing protein YAP associates with ErbB-4 and acts as a co-transcriptional activator for the carboxyl-terminal fragment of ErbB-4 that translocates to the nucleus
    • Komuro, A., Nagai, M., Navin, N. E. and Sudol, M. (2003) WW domain-containing protein YAP associates with ErbB-4 and acts as a co-transcriptional activator for the carboxyl-terminal fragment of ErbB-4 that translocates to the nucleus. J. Biol. Chem. 278, 33334-33341
    • (2003) J. Biol. Chem. , vol.278 , pp. 33334-33341
    • Komuro, A.1    Nagai, M.2    Navin, N.E.3    Sudol, M.4
  • 5
    • 0035873391 scopus 로고    scopus 로고
    • TEAD/TEF transcription factors utilize the activation domain of YAP65, a Src/Yes-associated protein localized in the cytoplasm
    • Vassilev, A., Kaneko, K. J., Shu, H., Zhao, Y. and DePamphilis, M. L. (2001) TEAD/TEF transcription factors utilize the activation domain of YAP65, a Src/Yes-associated protein localized in the cytoplasm. Genes Dev. 15, 1229-1241
    • (2001) Genes Dev. , vol.15 , pp. 1229-1241
    • Vassilev, A.1    Kaneko, K.J.2    Shu, H.3    Zhao, Y.4    DePamphilis, M.L.5
  • 6
    • 0033522472 scopus 로고    scopus 로고
    • A WW domain-containing Yes-associated protein (YAP) is a novel transcriptional co-activator
    • DOI 10.1093/emboj/18.9.2551
    • Yagi, R., Chen, L. F., Shigesada, K., Murakami, Y. and Ito, Y. (1999) A WW domain-containing yes-associated protein (YAP) is a novel transcriptional co-activator. EMBO J. 18, 2551-2562 (Pubitemid 29213286)
    • (1999) EMBO Journal , vol.18 , Issue.9 , pp. 2551-2562
    • Yagi, R.1    Chen, L.-F.2    Shigesada, K.3    Murakami, Y.4    Ito, Y.5
  • 7
    • 23744458034 scopus 로고    scopus 로고
    • The Hippo signaling pathway coordinately regulates cell proliferation and apoptosis by inactivating Yorkie, the Drosophila homolog of YAP
    • Huang, J., Wu, S., Barrera, J., Matthews, K. and Pan, D. (2005) The Hippo signaling pathway coordinately regulates cell proliferation and apoptosis by inactivating Yorkie, the Drosophila homolog of YAP. Cell 122, 421-434
    • (2005) Cell , vol.122 , pp. 421-434
    • Huang, J.1    Wu, S.2    Barrera, J.3    Matthews, K.4    Pan, D.5
  • 8
    • 0028997651 scopus 로고
    • Characterization of the mammalian YAP (Yes-associated protein) gene and its role in defining a novel protein module, the WW domain
    • Sudol, M., Bork, P., Einbond, A., Kastury, K., Druck, T., Negrini, M., Huebner, K. and Lehman, D. (1995) Characterization of the mammalian YAP (Yes-associated protein) gene and its role in defining a novel protein module, the WW domain. J. Biol. Chem. 270, 14733-14741
    • (1995) J. Biol. Chem. , vol.270 , pp. 14733-14741
    • Sudol, M.1    Bork, P.2    Einbond, A.3    Kastury, K.4    Druck, T.5    Negrini, M.6    Huebner, K.7    Lehman, D.8
  • 9
    • 0027988814 scopus 로고
    • The WW domain: A signalling site in dystrophin?
    • Bork, P. and Sudol, M. (1994) The WW domain: a signalling site in dystrophin? Trends Biochem. Sci. 19, 531-533
    • (1994) Trends Biochem. Sci. , vol.19 , pp. 531-533
    • Bork, P.1    Sudol, M.2
  • 10
    • 0029099161 scopus 로고
    • The WW domain of Yes-associated protein binds a proline-rich ligand that differs from the consensus established for Src homology 3-binding modules
    • Chen, H. I. and Sudol, M. (1995) The WW domain of Yes-associated protein binds a proline-rich ligand that differs from the consensus established for Src homology 3-binding modules. Proc. Natl. Acad. Sci. U.S.A. 92, 7819-7823
    • (1995) Proc. Natl. Acad. Sci. U.S.A. , vol.92 , pp. 7819-7823
    • Chen, H.I.1    Sudol, M.2
  • 11
    • 61749089153 scopus 로고    scopus 로고
    • The FERM-domain protein Expanded regulates Hippo pathway activity via direct interactions with the transcriptional activator Yorkie
    • Badouel, C., Gardano, L., Amin, N., Garg, A., Rosenfeld, R., Le Bihan, T. and McNeill, H. (2009) The FERM-domain protein Expanded regulates Hippo pathway activity via direct interactions with the transcriptional activator Yorkie. Dev. Cell 16, 411-420
    • (2009) Dev. Cell , vol.16 , pp. 411-420
    • Badouel, C.1    Gardano, L.2    Amin, N.3    Garg, A.4    Rosenfeld, R.5    Le Bihan, T.6    McNeill, H.7
  • 12
    • 0037249343 scopus 로고    scopus 로고
    • Akt phosphorylates the Yes-associated protein, YAP, to induce interaction with 14-3-3 and attenuation of p73-mediated apoptosis
    • Basu, S., Totty, N. F., Irwin, M. S., Sudol, M. and Downward, J. (2003) Akt phosphorylates the Yes-associated protein, YAP, to induce interaction with 14-3-3 and attenuation of p73-mediated apoptosis. Mol. Cell 11, 11-23
    • (2003) Mol. Cell , vol.11 , pp. 11-23
    • Basu, S.1    Totty, N.F.2    Irwin, M.S.3    Sudol, M.4    Downward, J.5
  • 13
    • 55549098921 scopus 로고    scopus 로고
    • Mst2 and Lats kinases regulate apoptotic function of Yes kinase-associated protein (YAP)
    • Oka, T., Mazack, V. and Sudol, M. (2008) Mst2 and Lats kinases regulate apoptotic function of Yes kinase-associated protein (YAP). J. Biol. Chem. 283, 27534-27546
    • (2008) J. Biol. Chem. , vol.283 , pp. 27534-27546
    • Oka, T.1    Mazack, V.2    Sudol, M.3
  • 14
    • 65449154339 scopus 로고    scopus 로고
    • Nuclear localization and pro-apoptotic signaling of YAP2 require intact PDZ-binding motif
    • Oka, T. and Sudol, M. (2009) Nuclear localization and pro-apoptotic signaling of YAP2 require intact PDZ-binding motif. Genes Cells 14, 607-615
    • (2009) Genes Cells , vol.14 , pp. 607-615
    • Oka, T.1    Sudol, M.2
  • 16
    • 0032941962 scopus 로고    scopus 로고
    • Differential behavior of E-cadherin and occludin in their colocalization with ZO-1 during the establishment of epithelial cell polarity
    • DOI 10.1002/(SICI)1097-4652(199905)179:2<115::AID-JCP1>3.0.CO;2-T
    • Ando-Akatsuka, Y., Yonemura, S., Itoh, M., Furuse, M. and Tsukita, S. (1999) Differential behavior of E-cadherin and occludin in their colocalization with ZO-1 during the establishment of epithelial cell polarity. J. Cell Physiol. 179, 115-125 (Pubitemid 29165139)
    • (1999) Journal of Cellular Physiology , vol.179 , Issue.2 , pp. 115-125
    • Ando-Akatsuka, Y.1    Yonemura, S.2    Itoh, M.3    Furuse, M.4    Tsukita, S.5
  • 17
    • 0032489875 scopus 로고    scopus 로고
    • ZO-3, a novel member of the MAGUK protein family found at the tight junction, interacts with ZO-1 and occludin
    • Haskins, J., Gu, L., Wittchen, E. S., Hibbard, J. and Stevenson, B. R. (1998) ZO-3, a novel member of the MAGUK protein family found at the tight junction, interacts with ZO-1 and occludin. J. Cell Biol. 141, 199-208
    • (1998) J. Cell Biol. , vol.141 , pp. 199-208
    • Haskins, J.1    Gu, L.2    Wittchen, E.S.3    Hibbard, J.4    Stevenson, B.R.5
  • 18
    • 0030748548 scopus 로고    scopus 로고
    • Involvement of ZO-1 in cadherin-based cell adhesion through its direct binding to α-catenin and actin filaments
    • Itoh, M., Nagafuchi, A., Moroi, S. and Tsukita, S. (1997) Involvement of ZO-1 in cadherin-based cell adhesion through its direct binding to α-catenin and actin filaments. J. Cell Biol. 138, 181-192
    • (1997) J. Cell Biol. , vol.138 , pp. 181-192
    • Itoh, M.1    Nagafuchi, A.2    Moroi, S.3    Tsukita, S.4
  • 19
    • 0033521140 scopus 로고    scopus 로고
    • Protein interactions at the tight junction. Actin has multiple binding partners, and ZO-1 forms independent complexes with ZO-2 and ZO-3
    • Wittchen, E. S., Haskins, J. and Stevenson, B. R. (1999) Protein interactions at the tight junction. Actin has multiple binding partners, and ZO-1 forms independent complexes with ZO-2 and ZO-3. J. Biol. Chem. 274, 35179-35185
    • (1999) J. Biol. Chem. , vol.274 , pp. 35179-35185
    • Wittchen, E.S.1    Haskins, J.2    Stevenson, B.R.3
  • 20
    • 0033825953 scopus 로고    scopus 로고
    • Molecular physiology and pathophysiology of tight junctions. I. Biogenesis of tight junctions and epithelial polarity
    • Cereijido, M., Shoshani, L. and Contreras, R. G. (2000) Molecular physiology and pathophysiology of tight junctions. I. Biogenesis of tight junctions and epithelial polarity. Am. J. Physiol. Gastrointest. Liver Physiol. 279, G477-G482
    • (2000) Am. J. Physiol. Gastrointest. Liver Physiol. , vol.279
    • Cereijido, M.1    Shoshani, L.2    Contreras, R.G.3
  • 22
    • 0032440448 scopus 로고    scopus 로고
    • The tight junction: Morphology to molecules
    • Stevenson, B. R. and Keon, B. H. (1998) The tight junction: morphology to molecules. Annu. Rev. Cell Dev. Biol. 14, 89-109
    • (1998) Annu. Rev. Cell Dev. Biol. , vol.14 , pp. 89-109
    • Stevenson, B.R.1    Keon, B.H.2
  • 23
    • 0032491391 scopus 로고    scopus 로고
    • The tight junction protein ZO-1 establishes a link between the transmembrane protein occludin and the actin cytoskeleton
    • Fanning, A. S., Jameson, B. J., Jesaitis, L. A. and Anderson, J. M. (1998) The tight junction protein ZO-1 establishes a link between the transmembrane protein occludin and the actin cytoskeleton. J. Biol. Chem. 273, 29745-29753
    • (1998) J. Biol. Chem. , vol.273 , pp. 29745-29753
    • Fanning, A.S.1    Jameson, B.J.2    Jesaitis, L.A.3    Anderson, J.M.4
  • 24
    • 0036832157 scopus 로고    scopus 로고
    • Isolation and functional characterization of the actin binding region in the tight junction protein ZO-1
    • Fanning, A. S., Ma, T. Y. and Anderson, J. M. (2002) Isolation and functional characterization of the actin binding region in the tight junction protein ZO-1. FASEB J. 16, 1835-1837
    • (2002) FASEB J. , vol.16 , pp. 1835-1837
    • Fanning, A.S.1    Ma, T.Y.2    Anderson, J.M.3
  • 25
    • 0034722381 scopus 로고    scopus 로고
    • Tight junction, a platform for trafficking and signaling protein complexes
    • Zahraoui, A., Louvard, D. and Galli, T. (2000) Tight junction, a platform for trafficking and signaling protein complexes. J. Cell Biol. 151, F31-36
    • (2000) J. Cell Biol. , vol.151
    • Zahraoui, A.1    Louvard, D.2    Galli, T.3
  • 26
    • 0029744106 scopus 로고    scopus 로고
    • The junction-associated protein, zonula occludens-1, localizes to the nucleus before the maturation and during the remodeling of cell-cell contacts
    • Gottardi, C. J., Arpin, M., Fanning, A. S. and Louvard, D. (1996) The junction-associated protein, zonula occludens-1, localizes to the nucleus before the maturation and during the remodeling of cell-cell contacts. Proc. Natl. Acad. Sci. U. S. A. 93, 10779-10784
    • (1996) Proc. Natl. Acad. Sci. U. S. A. , vol.93 , pp. 10779-10784
    • Gottardi, C.J.1    Arpin, M.2    Fanning, A.S.3    Louvard, D.4
  • 27
    • 0036343646 scopus 로고    scopus 로고
    • Nuclear localization of the tight junction protein ZO-2 in epithelial cells
    • Islas, S., Vega, J., Ponce, L. and Gonzalez-Mariscal, L. (2002) Nuclear localization of the tight junction protein ZO-2 in epithelial cells. Exp. Cell Res. 274, 138-148
    • (2002) Exp. Cell Res. , vol.274 , pp. 138-148
    • Islas, S.1    Vega, J.2    Ponce, L.3    Gonzalez-Mariscal, L.4
  • 28
    • 33748679558 scopus 로고    scopus 로고
    • The tight junction protein ZO-2 has several functional nuclear export signals
    • Gonzalez-Mariscal, L., Ponce, A., Alarcon, L. and Jaramillo, B. E. (2006) The tight junction protein ZO-2 has several functional nuclear export signals. Exp. Cell Res. 312, 3323-3335
    • (2006) Exp. Cell Res. , vol.312 , pp. 3323-3335
    • Gonzalez-Mariscal, L.1    Ponce, A.2    Alarcon, L.3    Jaramillo, B.E.4
  • 30
    • 0347986547 scopus 로고    scopus 로고
    • The tight junction protein ZO-2 associates with Jun, Fos and C/EBP transcription factors in epithelial cells
    • Betanzos, A., Huerta, M., Lopez-Bayghen, E., Azuara, E., Amerena, J. and Gonzalez-Mariscal, L. (2004) The tight junction protein ZO-2 associates with Jun, Fos and C/EBP transcription factors in epithelial cells. Exp. Cell Res. 292, 51-66
    • (2004) Exp. Cell Res. , vol.292 , pp. 51-66
    • Betanzos, A.1    Huerta, M.2    Lopez-Bayghen, E.3    Azuara, E.4    Amerena, J.5    Gonzalez-Mariscal, L.6
  • 31
    • 0034595219 scopus 로고    scopus 로고
    • The tight junction protein ZO-1 and an interacting transcription factor regulate ErbB-2 expression
    • Balda, M. S. and Matter, K. (2000) The tight junction protein ZO-1 and an interacting transcription factor regulate ErbB-2 expression. EMBO J. 19, 2024-2033
    • (2000) EMBO J. , vol.19 , pp. 2024-2033
    • Balda, M.S.1    Matter, K.2
  • 32
    • 33644773928 scopus 로고    scopus 로고
    • Regulation of PCNA and cyclin D1 expression and epithelial morphogenesis by the ZO-1-regulated transcription factor ZONAB/DbpA
    • Sourisseau, T., Georgiadis, A., Tsapara, A., Ali, R. R., Pestell, R., Matter, K. and Balda, M. S. (2006) Regulation of PCNA and cyclin D1 expression and epithelial morphogenesis by the ZO-1-regulated transcription factor ZONAB/DbpA. Mol. Cell. Biol. 26, 2387-2398
    • (2006) Mol. Cell. Biol. , vol.26 , pp. 2387-2398
    • Sourisseau, T.1    Georgiadis, A.2    Tsapara, A.3    Ali, R.R.4    Pestell, R.5    Matter, K.6    Balda, M.S.7
  • 33
    • 35948961118 scopus 로고    scopus 로고
    • Inactivation of YAP oncoprotein by the Hippo pathway is involved in cell contact inhibition and tissue growth control
    • Zhao, B., Wei, X., Li, W., Udan, R. S., Yang, Q., Kim, J., Xie, J., Ikenoue, T., Yu, J., Li, L. et al. (2007) Inactivation of YAP oncoprotein by the Hippo pathway is involved in cell contact inhibition and tissue growth control. Genes Dev. 21, 2747-2761
    • (2007) Genes Dev. , vol.21 , pp. 2747-2761
    • Zhao, B.1    Wei, X.2    Li, W.3    Udan, R.S.4    Yang, Q.5    Kim, J.6    Xie, J.7    Ikenoue, T.8    Yu, J.9    Li, L.10
  • 34
    • 34547643210 scopus 로고    scopus 로고
    • PDZ domains: Folding and binding
    • Jemth, P. and Gianni, S. (2007) PDZ domains: folding and binding. Biochemistry 46, 8701-8708
    • (2007) Biochemistry , vol.46 , pp. 8701-8708
    • Jemth, P.1    Gianni, S.2
  • 38
    • 0037462794 scopus 로고    scopus 로고
    • The tight junction protein ZO-2 localizes to the nucleus and interacts with the heterogeneous nuclear ribonucleoprotein scaffold attachment factor-B
    • Traweger, A., Fuchs, R., Krizbai, I. A., Weiger, T. M., Bauer, H. C. and Bauer, H. (2003) The tight junction protein ZO-2 localizes to the nucleus and interacts with the heterogeneous nuclear ribonucleoprotein scaffold attachment factor-B. J. Biol. Chem. 278, 2692-2700
    • (2003) J. Biol. Chem. , vol.278 , pp. 2692-2700
    • Traweger, A.1    Fuchs, R.2    Krizbai, I.A.3    Weiger, T.M.4    Bauer, H.C.5    Bauer, H.6
  • 39
    • 9444277252 scopus 로고    scopus 로고
    • Association of ARVCF with zonula occludens (ZO)-1 and ZO-2: Binding to PDZ-domain proteins and cell-cell adhesion regulate plasma membrane and nuclear localization of ARVCF
    • Kausalya, P. J., Phua, D. C. and Hunziker, W. (2004) Association of ARVCF with zonula occludens (ZO)-1 and ZO-2: binding to PDZ-domain proteins and cell-cell adhesion regulate plasma membrane and nuclear localization of ARVCF. Mol. Biol. Cell 15, 5503-5515
    • (2004) Mol. Biol. Cell , vol.15 , pp. 5503-5515
    • Kausalya, P.J.1    Phua, D.C.2    Hunziker, W.3
  • 41
    • 38049178969 scopus 로고    scopus 로고
    • Domain swapping within PDZ2 is responsible for dimerization of ZO proteins
    • Fanning, A. S., Lye, M. F., Anderson, J. M. and Lavie, A. (2007) Domain swapping within PDZ2 is responsible for dimerization of ZO proteins. J. Biol. Chem. 282, 37710-37716
    • (2007) J. Biol. Chem. , vol.282 , pp. 37710-37716
    • Fanning, A.S.1    Lye, M.F.2    Anderson, J.M.3    Lavie, A.4
  • 42
    • 33747686482 scopus 로고    scopus 로고
    • Dimerization of the scaffolding protein ZO-1 through the second PDZ domain
    • Utepbergenov, D. I., Fanning, A. S. and Anderson, J. M. (2006) Dimerization of the scaffolding protein ZO-1 through the second PDZ domain. J. Biol. Chem. 281, 24671-24677
    • (2006) J. Biol. Chem. , vol.281 , pp. 24671-24677
    • Utepbergenov, D.I.1    Fanning, A.S.2    Anderson, J.M.3
  • 44
    • 0033516532 scopus 로고    scopus 로고
    • Tight junction protein ZO-2 is differentially expressed in normal pancreatic ducts compared to human pancreatic adenocarcinoma
    • Chlenski, A., Ketels, K. V., Tsao, M. S., Talamonti, M. S., Anderson, M. R., Oyasu, R. and Scarpelli, D. G. (1999) Tight junction protein ZO-2 is differentially expressed in normal pancreatic ducts compared to human pancreatic adenocarcinoma. Int. J. Cancer 82, 137-144
    • (1999) Int. J. Cancer , vol.82 , pp. 137-144
    • Chlenski, A.1    Ketels, K.V.2    Tsao, M.S.3    Talamonti, M.S.4    Anderson, M.R.5    Oyasu, R.6    Scarpelli, D.G.7
  • 45
    • 42349084868 scopus 로고    scopus 로고
    • Negative regulation of YAP by LATS1 underscores evolutionary conservation of the Drosophila Hippo pathway
    • Zhang, J., Smolen, G. A. and Haber, D. A. (2008) Negative regulation of YAP by LATS1 underscores evolutionary conservation of the Drosophila Hippo pathway. Cancer Res. 68, 2789-2794
    • (2008) Cancer Res. , vol.68 , pp. 2789-2794
    • Zhang, J.1    Smolen, G.A.2    Haber, D.A.3
  • 46
    • 65249102236 scopus 로고    scopus 로고
    • YAP, TAZ, and Yorkie: A conserved family of signal-responsive transcriptional coregulators in animal development and human disease
    • Wang, K., Degerny, C., Xu, M. and Yang, X. J. (2009) YAP, TAZ, and Yorkie: a conserved family of signal-responsive transcriptional coregulators in animal development and human disease. Biochem. Cell Biol. 87, 77-91
    • (2009) Biochem. Cell Biol. , vol.87 , pp. 77-91
    • Wang, K.1    Degerny, C.2    Xu, M.3    Yang, X.J.4
  • 47
    • 78049290801 scopus 로고    scopus 로고
    • Modularity in the Hippo signaling pathway
    • doi: 10.1016/j.tibs.2010.05.010
    • Sudol, M. and Harvey, K. (2010) Modularity in the Hippo signaling pathway. Trends Biochem. Sci., doi: 10.1016/j.tibs.2010.05.010
    • (2010) Trends Biochem. Sci.
    • Sudol, M.1    Harvey, K.2
  • 50
    • 48749118705 scopus 로고    scopus 로고
    • The Fat and Warts signaling pathways: New insights into their regulation, mechanism and conservation
    • Reddy, B. V. and Irvine, K. D. (2008) The Fat and Warts signaling pathways: new insights into their regulation, mechanism and conservation. Development 135, 2827-2838
    • (2008) Development , vol.135 , pp. 2827-2838
    • Reddy, B.V.1    Irvine, K.D.2
  • 54
    • 70349308508 scopus 로고    scopus 로고
    • Cell regulation: Determined to signal discrete cooperation
    • Gibson, T. J. (2009) Cell regulation: determined to signal discrete cooperation. Trends Biochem. Sci. 34, 471-482
    • (2009) Trends Biochem. Sci. , vol.34 , pp. 471-482
    • Gibson, T.J.1


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