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Volumn 288, Issue 35, 2013, Pages 25244-25253

Regulation of hypoxia-inducible factor 1α (HIF-1α) by lysophosphatidic acid is dependent on interplay between p53 and Krüppel-like factor 5

Author keywords

[No Author keywords available]

Indexed keywords

COLON CANCER; HYPOXIA-INDUCIBLE FACTOR 1; LYSOPHOSPHATIDIC ACID; NEW DIMENSIONS;

EID: 84883313474     PISSN: 00219258     EISSN: 1083351X     Source Type: Journal    
DOI: 10.1074/jbc.M113.489708     Document Type: Article
Times cited : (60)

References (55)
  • 1
    • 84880920150 scopus 로고    scopus 로고
    • Clinicopathological significance of hypoxia-inducible factor- 1 alpha (HIF-1α) expression in gastric cancer
    • Isobe, T., Aoyagi, K., Koufuji, K., Shirouzu, K., Kawahara, A., Taira, T., and Kage, M. (2013) Clinicopathological significance of hypoxia-inducible factor- 1 alpha (HIF-1α) expression in gastric cancer. Int. J. Clin. Oncol. 18, 293-304
    • (2013) Int. J. Clin. Oncol , vol.18 , pp. 293-304
    • Isobe, T.1    Aoyagi, K.2    Koufuji, K.3    Shirouzu, K.4    Kawahara, A.5    Taira, T.6    Kage, M.7
  • 3
    • 84856739946 scopus 로고    scopus 로고
    • Hypoxia-inducible factors in physiology and medicine
    • Semenza, G. L. (2012) Hypoxia-inducible factors in physiology and medicine. Cell 148, 399-408
    • (2012) Cell , vol.148 , pp. 399-408
    • Semenza, G.L.1
  • 5
    • 0036260697 scopus 로고    scopus 로고
    • Cellular response to oxidative stress: Signaling for suicide and survival
    • Martindale, J. L., and Holbrook, N. J. (2002) Cellular response to oxidative stress: signaling for suicide and survival. J. Cell Physiol. 192, 1-15
    • (2002) J. Cell Physiol , vol.192 , pp. 1-15
    • Martindale, J.L.1    Holbrook, N.J.2
  • 6
    • 77956713136 scopus 로고    scopus 로고
    • Hypoxia-inducible factor-1 activation in nonhypoxic conditions: The essential role of mitochondrial-derived reactive oxygen species
    • Patten, D. A., Lafleur, V. N., Robitaille, G. A., Chan, D. A., Giaccia, A. J., and Richard, D. E. (2010) Hypoxia-inducible factor-1 activation in nonhypoxic conditions: the essential role of mitochondrial-derived reactive oxygen species. Mol. Biol. Cell 21, 3247-3257
    • (2010) Mol. Biol. Cell , vol.21 , pp. 3247-3257
    • Patten, D.A.1    Lafleur, V.N.2    Robitaille, G.A.3    Chan, D.A.4    Giaccia, A.J.5    Richard, D.E.6
  • 7
    • 32944477859 scopus 로고    scopus 로고
    • Hypoxia-inducible factor-1α promotes nonhypoxia-mediated proliferation in colon cancer cells and xenografts
    • Dang, D. T., Chen, F., Gardner, L. B., Cummins, J. M., Rago, C., Bunz, F., Kantsevoy, S. V., and Dang, L. H. (2006) Hypoxia-inducible factor-1α promotes nonhypoxia-mediated proliferation in colon cancer cells and xenografts. Cancer Res. 66, 1684-1936
    • (2006) Cancer Res , vol.66 , pp. 1684-1936
    • Dang, D.T.1    Chen, F.2    Gardner, L.B.3    Cummins, J.M.4    Rago, C.5    Bunz, F.6    Kantsevoy, S.V.7    Dang, L.H.8
  • 9
    • 4344706187 scopus 로고    scopus 로고
    • The ins and outs of lysophosphatidic acid signaling
    • Moolenaar, W. H., van Meeteren, L. A., and Giepmans, B. N. (2004) The ins and outs of lysophosphatidic acid signaling. Bioessays 26, 870-881
    • (2004) Bioessays , vol.26 , pp. 870-881
    • Moolenaar, W.H.1    Van Meeteren, L.A.2    Giepmans, B.N.3
  • 11
    • 79952292419 scopus 로고    scopus 로고
    • MAGI-3 competes with NHERF-2 to negatively regulate LPA2 receptor signaling in colon cancer cells
    • Lee, S. J., Ritter, S. L., Zhang, H., Shim, H., Hall, R. A., and Yun, C. C. (2011) MAGI-3 competes with NHERF-2 to negatively regulate LPA2 receptor signaling in colon cancer cells. Gastroenterology 140, 923-934
    • (2011) Gastroenterology , vol.140 , pp. 923-934
    • Lee, S.J.1    Ritter, S.L.2    Zhang, H.3    Shim, H.4    Hall, R.A.5    Yun, C.C.6
  • 12
    • 84878971521 scopus 로고    scopus 로고
    • Distinct phospholipase C-β isozymes mediate lysophosphatidic acid receptor 1 effects on intestinal epithelial homeostasis and wound closure
    • Lee, S. J., Leoni, G., Neumann, P. A., Chun, J., Nusrat, A., and Yun, C. C. (2013) Distinct phospholipase C-β isozymes mediate lysophosphatidic acid receptor 1 effects on intestinal epithelial homeostasis and wound closure. Mol. Cell. Biol. 33, 2016-2028
    • (2013) Mol. Cell. Biol , vol.33 , pp. 2016-2028
    • Lee, S.J.1    Leoni, G.2    Neumann, P.A.3    Chun, J.4    Nusrat, A.5    Yun, C.C.6
  • 15
    • 77956034731 scopus 로고    scopus 로고
    • Aiming drug discovery at lysophosphatidic acid targets
    • Tigyi, G. (2010) Aiming drug discovery at lysophosphatidic acid targets. Br. J. Pharmacol. 161, 241-270
    • (2010) Br. J. Pharmacol , vol.161 , pp. 241-270
    • Tigyi, G.1
  • 16
    • 78149481269 scopus 로고    scopus 로고
    • The absence of LPA receptor 2 reduces the tumorigenesis by ApcMin mutation in the intestine
    • Lin, S., Lee, S. J., Shim, H., Chun, J., and Yun, C. C. (2010) The absence of LPA receptor 2 reduces the tumorigenesis by ApcMin mutation in the intestine. Am. J. Physiol. Gastrointest. Liver Physiol. 299, G1128-G1138
    • (2010) Am. J. Physiol. Gastrointest. Liver Physiol , vol.299
    • Lin, S.1    Lee, S.J.2    Shim, H.3    Chun, J.4    Yun, C.C.5
  • 17
    • 65249173527 scopus 로고    scopus 로고
    • The absence of LPA2 attenuates tumor formation in an experimental model of colitis-associated cancer
    • Lin, S., Wang, D., Iyer, S., Ghaleb, A. M., Shim, H., Yang, V. W., Chun, J., and Yun, C. C. (2009) The absence of LPA2 attenuates tumor formation in an experimental model of colitis-associated cancer. Gastroenterology 136, 1711-1720
    • (2009) Gastroenterology , vol.136 , pp. 1711-1720
    • Lin, S.1    Wang, D.2    Iyer, S.3    Ghaleb, A.M.4    Shim, H.5    Yang, V.W.6    Chun, J.7    Yun, C.C.8
  • 18
    • 0035897531 scopus 로고    scopus 로고
    • Lysophosphatidic acid induction of vascular endothelial growth factor expression in human ovarian cancer cells
    • Hu, Y. L., Tee, M. K., Goetzl, E. J., Auersperg, N., Mills, G. B., Ferrara, N., and Jaffe, R. B. (2001) Lysophosphatidic acid induction of vascular endothelial growth factor expression in human ovarian cancer cells. J. Natl. Cancer Inst. 93, 762-768
    • (2001) J. Natl. Cancer Inst , vol.93 , pp. 762-768
    • Hu, Y.L.1    Tee, M.K.2    Goetzl, E.J.3    Auersperg, N.4    Mills, G.B.5    Ferrara, N.6    Jaffe, R.B.7
  • 19
    • 0942301169 scopus 로고    scopus 로고
    • Induction of protein growth factor systems in the ovaries of transgenic mice overexpressing human type 2 lysophosphatidic acid G protein-coupled receptor (LPA2)
    • Huang, M. C., Lee, H. Y., Yeh, C. C., Kong, Y., Zaloudek, C. J., and Goetzl, E. J. (2004) Induction of protein growth factor systems in the ovaries of transgenic mice overexpressing human type 2 lysophosphatidic acid G protein-coupled receptor (LPA2). Oncogene 23, 122-129
    • (2004) Oncogene , vol.23 , pp. 122-129
    • Huang, M.C.1    Lee, H.Y.2    Yeh, C.C.3    Kong, Y.4    Zaloudek, C.J.5    Goetzl, E.J.6
  • 20
    • 33748056087 scopus 로고    scopus 로고
    • Hypoxia enhances lysophosphatidic acid responsiveness in ovarian cancer cells and lysophosphatidic acid induces ovarian tumor metastasis in vivo
    • Kim, K. S., Sengupta, S., Berk, M., Kwak, Y. G., Escobar, P. F., Belinson, J., Mok, S. C., and Xu, Y. (2006) Hypoxia enhances lysophosphatidic acid responsiveness in ovarian cancer cells and lysophosphatidic acid induces ovarian tumor metastasis in vivo. Cancer Res. 66, 7983-7990
    • (2006) Cancer Res , vol.66 , pp. 7983-7990
    • Kim, K.S.1    Sengupta, S.2    Berk, M.3    Kwak, Y.G.4    Escobar, P.F.5    Belinson, J.6    Mok, S.C.7    Xu, Y.8
  • 23
    • 34447522119 scopus 로고    scopus 로고
    • Lysophosphatidic acid facilitates proliferation of colon cancer cells via induction of Krüppel-like factor 5
    • Zhang, H., Bialkowska, A., Rusovici, R., Chanchevalap, S., Shim, H., Katz, J. P., Yang, V. W., and Yun, C. C. (2007) Lysophosphatidic acid facilitates proliferation of colon cancer cells via induction of Krüppel-like factor 5. J. Biol. Chem. 282, 15541-15549
    • (2007) J. Biol. Chem , vol.282 , pp. 15541-15549
    • Zhang, H.1    Bialkowska, A.2    Rusovici, R.3    Chanchevalap, S.4    Shim, H.5    Katz, J.P.6    Yang, V.W.7    Yun, C.C.8
  • 24
    • 43249104714 scopus 로고    scopus 로고
    • Different mutant/wild-type p53 combinations cause a spectrum of increased invasive potential in nonmalignant immortalized human mammary epithelial cells
    • Junk, D. J., Vrba, L., Watts, G. S., Oshiro, M. M., Martinez, J. D., and Futscher, B. W. (2008) Different mutant/wild-type p53 combinations cause a spectrum of increased invasive potential in nonmalignant immortalized human mammary epithelial cells. Neoplasia 10, 450-461
    • (2008) Neoplasia , vol.10 , pp. 450-461
    • Junk, D.J.1    Vrba, L.2    Watts, G.S.3    Oshiro, M.M.4    Martinez, J.D.5    Futscher, B.W.6
  • 26
    • 0036190875 scopus 로고    scopus 로고
    • PROMO: Detection of known transcription regulatory elements using species-tailored searches
    • Messeguer, X., Escudero, R., Farré, D., Núñez, O., Martínez, J., and Albà, M. M. (2002) PROMO: detection of known transcription regulatory elements using species-tailored searches. Bioinformatics 18, 333-334
    • (2002) Bioinformatics , vol.18 , pp. 333-334
    • Messeguer, X.1    Escudero, R.2    Farré, D.3    Núñez, O.4    Martínez, J.5    Albà, M.M.6
  • 27
    • 80054700545 scopus 로고    scopus 로고
    • Serum- and glucocorticoid-induced kinase 3 in recycling endosomes mediates acute activation of Na+/H+ exchanger NHE3 by glucocorticoids
    • He, P., Lee, S. J., Lin, S., Seidler, U., Lang, F., Fejes-Toth, G., Naray-Fejes-Toth, A., and Yun, C. C. (2011) Serum- and glucocorticoid-induced kinase 3 in recycling endosomes mediates acute activation of Na+/H+ exchanger NHE3 by glucocorticoids. Mol. Biol. Cell 22, 3812-3825
    • (2011) Mol. Biol. Cell , vol.22 , pp. 3812-3825
    • He, P.1    Lee, S.J.2    Lin, S.3    Seidler, U.4    Lang, F.5    Fejes-Toth, G.6    Naray-Fejes-Toth, A.7    Yun, C.C.8
  • 29
    • 36248947897 scopus 로고    scopus 로고
    • Lysophosphatidic acid decreases the nuclear localization and cellular abundance of the p53 tumor suppressor in A549 lung carcinoma cells
    • Murph, M. M., Hurst-Kennedy, J., Newton, V., Brindley, D. N., and Radhakrishna, H. (2007) Lysophosphatidic acid decreases the nuclear localization and cellular abundance of the p53 tumor suppressor in A549 lung carcinoma cells. Mol. Cancer Res. 5, 1201-1211
    • (2007) Mol. Cancer Res , vol.5 , pp. 1201-1211
    • Murph, M.M.1    Hurst-Kennedy, J.2    Newton, V.3    Brindley, D.N.4    Radhakrishna, H.5
  • 31
    • 77957284773 scopus 로고    scopus 로고
    • 24R,25-Dihydroxyvitamin D3, lysophosphatidic acid, and p53: A signaling axis in the inhibition of phosphate-induced chondrocyte apoptosis
    • Hurst-Kennedy, J., Zhong, M., Gupta, V., Boyan, B. D., and Schwartz, Z. (2010) 24R,25-Dihydroxyvitamin D3, lysophosphatidic acid, and p53: a signaling axis in the inhibition of phosphate-induced chondrocyte apoptosis. J. Steroid Biochem. Mol. Biol. 122, 264-271
    • (2010) J. Steroid Biochem. Mol. Biol , vol.122 , pp. 264-271
    • Hurst-Kennedy, J.1    Zhong, M.2    Gupta, V.3    Boyan, B.D.4    Schwartz, Z.5
  • 32
    • 0025312728 scopus 로고
    • A genetic model for colorectal tumorigenesis
    • Fearon, E. R., and Vogelstein, B. (1990) A genetic model for colorectal tumorigenesis. Cell 61, 759-767
    • (1990) Cell , vol.61 , pp. 759-767
    • Fearon, E.R.1    Vogelstein, B.2
  • 34
    • 17844382139 scopus 로고    scopus 로고
    • Gprotein-coupled lysophosphatidic acid receptors stimulate proliferation of colon cancer cells through the β-catenin pathway
    • Yang, M., Zhong, W. W., Srivastava, N., Slavin, A., Yang, J., Hoey, T., and An, S. (2005)Gprotein-coupled lysophosphatidic acid receptors stimulate proliferation of colon cancer cells through the β-catenin pathway. Proc. Natl. Acad. Sci. U.S.A. 102, 6027-6032
    • (2005) Proc. Natl. Acad. Sci. U.S.A. , vol.102 , pp. 6027-6032
    • Yang, M.1    Zhong, W.W.2    Srivastava, N.3    Slavin, A.4    Yang, J.5    Hoey, T.6    An, S.7
  • 35
    • 0023339504 scopus 로고
    • Molecular analysis and chromosomal mapping of amplified genes isolated from a transformed mouse 3T3 cell line
    • Cahilly-Snyder, L., Yang-Feng, T., Francke, U., and George, D. L. (1987) Molecular analysis and chromosomal mapping of amplified genes isolated from a transformed mouse 3T3 cell line. Somat. Cell Mol. Genet. 13, 235-244
    • (1987) Somat. Cell Mol. Genet , vol.13 , pp. 235-244
    • Cahilly-Snyder, L.1    Yang-Feng, T.2    Francke, U.3    George, D.L.4
  • 36
    • 0026649648 scopus 로고
    • The mdm-2 oncogene product forms a complex with the p53 protein and inhibits p53-mediated transactivation
    • Momand, J., Zambetti, G. P., Olson, D. C., George, D., and Levine, A. J. (1992) The mdm-2 oncogene product forms a complex with the p53 protein and inhibits p53-mediated transactivation. Cell 69, 1237-1245
    • (1992) Cell , vol.69 , pp. 1237-1245
    • Momand, J.1    Zambetti, G.P.2    Olson, D.C.3    George, D.4    Levine, A.J.5
  • 37
    • 77955871461 scopus 로고    scopus 로고
    • The Mdm2-p53 relationship evolves: Mdm2 swings both ways as an oncogene and a tumor suppressor
    • Manfredi, J. J. (2010) The Mdm2-p53 relationship evolves: Mdm2 swings both ways as an oncogene and a tumor suppressor. Genes Dev. 24, 1580-1589
    • (2010) Genes Dev , vol.24 , pp. 1580-1589
    • Manfredi, J.J.1
  • 38
    • 0034735904 scopus 로고    scopus 로고
    • MDM2-dependent ubiquitination of nuclear and cytoplasmic P53
    • Yu, Z. K., Geyer, R. K., and Maki, C. G. (2000) MDM2-dependent ubiquitination of nuclear and cytoplasmic P53. Oncogene 19, 5892-5897
    • (2000) Oncogene , vol.19 , pp. 5892-5897
    • Yu, Z.K.1    Geyer, R.K.2    Maki, C.G.3
  • 39
    • 0029131447 scopus 로고
    • A functional p53-responsive intronic promoter is contained within the human mdm2 gene
    • Zauberman, A., Flusberg, D., Haupt, Y., Barak, Y., and Oren, M. (1995) A functional p53-responsive intronic promoter is contained within the human mdm2 gene. Nucleic Acids Res. 23, 2584-2592
    • (1995) Nucleic Acids Res , vol.23 , pp. 2584-2592
    • Zauberman, A.1    Flusberg, D.2    Haupt, Y.3    Barak, Y.4    Oren, M.5
  • 40
    • 0027459198 scopus 로고
    • Mdm2 expression is induced by wild type p53 activity
    • Barak, Y., Juven, T., Haffner, R., and Oren, M. (1993) mdm2 expression is induced by wild type p53 activity. EMBO J. 12, 461-468
    • (1993) EMBO J , vol.12 , pp. 461-468
    • Barak, Y.1    Juven, T.2    Haffner, R.3    Oren, M.4
  • 41
    • 33751297349 scopus 로고    scopus 로고
    • Activation of hypoxia-inducible factor-1α is necessary for lysophosphatidic acid-induced vascular endothelial growth factor expression
    • Lee, J., Park, S. Y., Lee, E. K., Park, C. G., Chung, H. C., Rha, S. Y., Kim, Y. K., Bae, G. U., Kim, B. K., Han, J. W., and Lee, H. Y. (2006) Activation of hypoxia-inducible factor-1α is necessary for lysophosphatidic acid-induced vascular endothelial growth factor expression. Clin. Cancer Res. 12, 6351-6358
    • (2006) Clin. Cancer Res , vol.12 , pp. 6351-6358
    • Lee, J.1    Park, S.Y.2    Lee, E.K.3    Park, C.G.4    Chung, H.C.5    Rha, S.Y.6    Kim, Y.K.7    Bae, G.U.8    Kim, B.K.9    Han, J.W.10    Lee, H.Y.11
  • 42
    • 0037079250 scopus 로고    scopus 로고
    • P53 mutation as a source of aberrant β-catenin accumulation in cancer cells
    • Cagatay, T., and Ozturk, M. (2002) P53 mutation as a source of aberrant β-catenin accumulation in cancer cells. Oncogene 21, 7971-7980
    • (2002) Oncogene , vol.21 , pp. 7971-7980
    • Cagatay, T.1    Ozturk, M.2
  • 43
    • 0026760566 scopus 로고
    • Abnormal structure and expression of the p53 gene in human ovarian carcinoma cell lines
    • Yaginuma, Y., and Westphal, H. (1992) Abnormal structure and expression of the p53 gene in human ovarian carcinoma cell lines. Cancer Res. 52, 4196-4199
    • (1992) Cancer Res , vol.52 , pp. 4196-4199
    • Yaginuma, Y.1    Westphal, H.2
  • 46
    • 0033104158 scopus 로고    scopus 로고
    • A gene encoding an intestinal-enriched member of the Krüppel-like factor family expressed in intestinal epithelial cells
    • Conkright, M. D., Wani, M. A., Anderson, K. P., and Lingrel, J. B. (1999) A gene encoding an intestinal-enriched member of the Krüppel-like factor family expressed in intestinal epithelial cells. Nucleic Acids Res. 27, 1263-1270
    • (1999) Nucleic Acids Res , vol.27 , pp. 1263-1270
    • Conkright, M.D.1    Wani, M.A.2    Anderson, K.P.3    Lingrel, J.B.4
  • 47
    • 34250216304 scopus 로고    scopus 로고
    • The diverse functions of Krüppel-like factors 4 and 5 in epithelial biology and pathobiology
    • McConnell, B. B., Ghaleb, A. M., Nandan, M. O., and Yang, V. W. (2007) The diverse functions of Krüppel-like factors 4 and 5 in epithelial biology and pathobiology. Bioessays 29, 549-557
    • (2007) Bioessays , vol.29 , pp. 549-557
    • McConnell, B.B.1    Ghaleb, A.M.2    Nandan, M.O.3    Yang, V.W.4
  • 48
    • 2442686555 scopus 로고    scopus 로고
    • Krüppel-like factor 5 mediates the transforming activity of oncogenic H-Ras
    • Nandan, M. O., Yoon, H. S., Zhao, W., Ouko, L. A., Chanchevalap, S., and Yang, V. W. (2004) Krüppel-like factor 5 mediates the transforming activity of oncogenic H-Ras. Oncogene 23, 3404-3413
    • (2004) Oncogene , vol.23 , pp. 3404-3413
    • Nandan, M.O.1    Yoon, H.S.2    Zhao, W.3    Ouko, L.A.4    Chanchevalap, S.5    Yang, V.W.6
  • 50
    • 33744955883 scopus 로고    scopus 로고
    • KLF5 Interacts with p53 in regulating survivin expression in acute lymphoblastic leukemia
    • Zhu, N., Gu, L., Findley, H. W., Chen, C., Dong, J. T., Yang, L., and Zhou, M. (2006) KLF5 Interacts with p53 in regulating survivin expression in acute lymphoblastic leukemia. J. Biol. Chem. 281, 14711-14718
    • (2006) J. Biol. Chem , vol.281 , pp. 14711-14718
    • Zhu, N.1    Gu, L.2    Findley, H.W.3    Chen, C.4    Dong, J.T.5    Yang, L.6    Zhou, M.7
  • 51
    • 80054015502 scopus 로고    scopus 로고
    • Loss of transcription factor KLF5 in the context of p53 ablation drives invasive progression of human squamous cell cancer
    • Yang, Y., Nakagawa, H., Tetreault, M. P., Billig, J., Victor, N., Goyal, A., Sepulveda, A. R., and Katz, J. P. (2011) Loss of transcription factor KLF5 in the context of p53 ablation drives invasive progression of human squamous cell cancer. Cancer Res. 71, 6475-6484
    • (2011) Cancer Res , vol.71 , pp. 6475-6484
    • Yang, Y.1    Nakagawa, H.2    Tetreault, M.P.3    Billig, J.4    Victor, N.5    Goyal, A.6    Sepulveda, A.R.7    Katz, J.P.8
  • 53
    • 0028897398 scopus 로고
    • Frequent and characteristic K-ras activation and absence of p53 protein accumulation in aberrant crypt foci of the colon
    • Yamashita, N., Minamoto, T., Ochiai, A., Onda, M., and Esumi, H. (1995) Frequent and characteristic K-ras activation and absence of p53 protein accumulation in aberrant crypt foci of the colon. Gastroenterology 108, 434-440
    • (1995) Gastroenterology , vol.108 , pp. 434-440
    • Yamashita, N.1    Minamoto, T.2    Ochiai, A.3    Onda, M.4    Esumi, H.5
  • 54
    • 34347240776 scopus 로고    scopus 로고
    • Lysophosphatidic acid prevents apoptosis of Caco-2 colon cancer cells via activation of mitogen-activated protein kinase and phosphorylation of Bad
    • Rusovici, R., Ghaleb, A., Shim, H., Yang, V. W., and Yun, C. C. (2007) Lysophosphatidic acid prevents apoptosis of Caco-2 colon cancer cells via activation of mitogen-activated protein kinase and phosphorylation of Bad. Biochim. Biophys. Acta 1770, 1194-1203
    • (2007) Biochim. Biophys. Acta , vol.1770 , pp. 1194-1203
    • Rusovici, R.1    Ghaleb, A.2    Shim, H.3    Yang, V.W.4    Yun, C.C.5


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