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Volumn 8, Issue 7, 2013, Pages

The Poplar MYB Master Switches Bind to the SMRE Site and Activate the Secondary Wall Biosynthetic Program during Wood Formation

Author keywords

[No Author keywords available]

Indexed keywords

PROTEIN EGMYB2; PROTEIN MYB; PROTEIN MYB46; PROTEIN MYB83; PROTEIN PTMYB4; PROTEIN PTRMYB2; PROTEIN PTRMYB20; PROTEIN PTRMYB21; PROTEIN PTRMYB3; UNCLASSIFIED DRUG;

EID: 84880804847     PISSN: None     EISSN: 19326203     Source Type: Journal    
DOI: 10.1371/journal.pone.0069219     Document Type: Article
Times cited : (118)

References (50)
  • 2
    • 24044441630 scopus 로고    scopus 로고
    • Transcriptome profiling of vertical stem segments provides insights into the genetic regulation of secondary growth in hybrid aspen trees
    • Prassinos C, Ko JH, Yang J, Han KH, (2005) Transcriptome profiling of vertical stem segments provides insights into the genetic regulation of secondary growth in hybrid aspen trees. Plant Cell Physiol 46: 1213-1225.
    • (2005) Plant Cell Physiol , vol.46 , pp. 1213-1225
    • Prassinos, C.1    Ko, J.H.2    Yang, J.3    Han, K.H.4
  • 3
    • 33644924586 scopus 로고    scopus 로고
    • Biosynthesis of cellulose-enriched tension wood in Populus: global analysis of transcripts and metabolites identifies biochemical and developmental regulators in secondary wall biosynthesis
    • Andersson-Gunneras S, Mellerowicz EJ, Love J, Segerman B, Ohmiya Y, et al. (2006) Biosynthesis of cellulose-enriched tension wood in Populus: global analysis of transcripts and metabolites identifies biochemical and developmental regulators in secondary wall biosynthesis. Plant J 45: 144-165.
    • (2006) Plant J , vol.45 , pp. 144-165
    • Andersson-Gunneras, S.1    Mellerowicz, E.J.2    Love, J.3    Segerman, B.4    Ohmiya, Y.5
  • 4
    • 55649115573 scopus 로고    scopus 로고
    • Identification of conserved core xylem gene sets: conifer cDNA microarray development, transcript profiling and computational analyses
    • Pavy N, Boyle B, Nelson C, Paule C, Giguère I, et al. (2008) Identification of conserved core xylem gene sets: conifer cDNA microarray development, transcript profiling and computational analyses. New Phytol 180: 766-786.
    • (2008) New Phytol , vol.180 , pp. 766-786
    • Pavy, N.1    Boyle, B.2    Nelson, C.3    Paule, C.4    Giguère, I.5
  • 5
    • 60249102826 scopus 로고    scopus 로고
    • Expansion and diversification of the Populus R2R3-MYB family of transcription factors
    • Wilkins O, Nahal H, Foong J, Provart NJ, Campbell MM, (2009) Expansion and diversification of the Populus R2R3-MYB family of transcription factors. Plant Physiol 149: 981-993.
    • (2009) Plant Physiol , vol.149 , pp. 981-993
    • Wilkins, O.1    Nahal, H.2    Foong, J.3    Provart, N.J.4    Campbell, M.M.5
  • 6
    • 84907954471 scopus 로고    scopus 로고
    • Genome-wide transcriptome analysis of the transition from primary to secondary stem development in Populus trichocarpa
    • Dharmawardhana P, Brunner AM, Strauss SH, (2010) Genome-wide transcriptome analysis of the transition from primary to secondary stem development in Populus trichocarpa. BMC Genomics 11: 150.
    • (2010) BMC Genomics , vol.11 , pp. 150
    • Dharmawardhana, P.1    Brunner, A.M.2    Strauss, S.H.3
  • 8
    • 4544352472 scopus 로고    scopus 로고
    • A high-resolution transcript profile across the wood-forming meristem of poplar identifies potential regulators of cambial stem cell identity
    • Schrader J, Nilsson J, Mellerowicz E, Berglund A, Nilsson P, et al. (2004) A high-resolution transcript profile across the wood-forming meristem of poplar identifies potential regulators of cambial stem cell identity. Plant Cell 16: 2278-2292.
    • (2004) Plant Cell , vol.16 , pp. 2278-2292
    • Schrader, J.1    Nilsson, J.2    Mellerowicz, E.3    Berglund, A.4    Nilsson, P.5
  • 9
    • 71649100687 scopus 로고    scopus 로고
    • The Populus homeobox gene ARBORKNOX2 regulates cell differentiation during secondary growth
    • Du J, Mansfield SD, Groover AT, (2009) The Populus homeobox gene ARBORKNOX2 regulates cell differentiation during secondary growth. Plant J 60: 1000-1014.
    • (2009) Plant J , vol.60 , pp. 1000-1014
    • Du, J.1    Mansfield, S.D.2    Groover, A.T.3
  • 10
    • 79952215358 scopus 로고    scopus 로고
    • The Populus Class III HD ZIP transcription factor POPCORONA affects cell differentiation during secondary growth of woody stems
    • Du J, Miura E, Robischon M, Martinez C, Groover A, (2011) The Populus Class III HD ZIP transcription factor POPCORONA affects cell differentiation during secondary growth of woody stems. PLoS One 6: e17458.
    • (2011) PLoS One , vol.6
    • Du, J.1    Miura, E.2    Robischon, M.3    Martinez, C.4    Groover, A.5
  • 11
    • 79952290449 scopus 로고    scopus 로고
    • The Populus class III HD ZIP, popREVOLUTA, influences cambium initiation and patterning of woody stems
    • Robischon M, Du J, Miura E, Groover A, (2011) The Populus class III HD ZIP, popREVOLUTA, influences cambium initiation and patterning of woody stems. Plant Physiol 155: 1214-1225.
    • (2011) Plant Physiol , vol.155 , pp. 1214-1225
    • Robischon, M.1    Du, J.2    Miura, E.3    Groover, A.4
  • 13
    • 78049463790 scopus 로고    scopus 로고
    • Evolutionary conservation of the transcriptional network regulating secondary cell wall biosynthesis
    • Zhong R, Lee C, Ye Z-H, (2010) Evolutionary conservation of the transcriptional network regulating secondary cell wall biosynthesis. Trends Plant Sci 15: 625-631.
    • (2010) Trends Plant Sci , vol.15 , pp. 625-631
    • Zhong, R.1    Lee, C.2    Ye, Z.-H.3
  • 14
    • 75949130469 scopus 로고    scopus 로고
    • Functional characterization of poplar wood-associated NAC domain transcription factors
    • Zhong R, Lee C, Ye ZH, (2010) Functional characterization of poplar wood-associated NAC domain transcription factors. Plant Physiol 152: 1044-1055.
    • (2010) Plant Physiol , vol.152 , pp. 1044-1055
    • Zhong, R.1    Lee, C.2    Ye, Z.H.3
  • 15
    • 80455173514 scopus 로고    scopus 로고
    • Dissection of the transcriptional program regulating secondary wall biosynthesis during wood formation in poplar
    • Zhong R, McCarthy RL, Lee C, Ye Z-H, (2011) Dissection of the transcriptional program regulating secondary wall biosynthesis during wood formation in poplar. Plant Physiol 157: 1452-1468.
    • (2011) Plant Physiol , vol.157 , pp. 1452-1468
    • Zhong, R.1    McCarthy, R.L.2    Lee, C.3    Ye, Z.-H.4
  • 16
    • 77955007863 scopus 로고    scopus 로고
    • The poplar PtrWNDs are transcriptional activators of secondary cell wall biosynthesis
    • Zhong R, Ye Z-H, (2010) The poplar PtrWNDs are transcriptional activators of secondary cell wall biosynthesis. Plant Signal Behav 5: 469-472.
    • (2010) Plant Signal Behav , vol.5 , pp. 469-472
    • Zhong, R.1    Ye, Z.-H.2
  • 17
    • 79960918743 scopus 로고    scopus 로고
    • A NAC domain protein family contributing to the regulation of wood formation in poplar. Plant J
    • Ohtani M, Nishikubo N, Xu B, Yamaguchi M, Mitsuda N, et al. (2011) A NAC domain protein family contributing to the regulation of wood formation in poplar. Plant J. 67: 499-512.
    • (2011) , vol.67 , pp. 499-512
    • Ohtani, M.1    Nishikubo, N.2    Xu, B.3    Yamaguchi, M.4    Mitsuda, N.5
  • 18
    • 77954291233 scopus 로고    scopus 로고
    • An NAC transcription factor orchestrates multiple features of cell wall development in Medicago truncatula
    • Zhao Q, Gallego-Giraldo L, Wang H, Zeng Y, Ding SY, et al. (2010) An NAC transcription factor orchestrates multiple features of cell wall development in Medicago truncatula. Plant J 63: 100-114.
    • (2010) Plant J , vol.63 , pp. 100-114
    • Zhao, Q.1    Gallego-Giraldo, L.2    Wang, H.3    Zeng, Y.4    Ding, S.Y.5
  • 19
    • 80054685612 scopus 로고    scopus 로고
    • Transcriptional activation of secondary wall biosynthesis by rice and maize NAC and MYB transcription factors
    • Zhong R, Lee C, McCarthy RL, Reeves CK, Jones EG, et al. (2011) Transcriptional activation of secondary wall biosynthesis by rice and maize NAC and MYB transcription factors. Plant Cell Physiol 52: 1856-1871.
    • (2011) Plant Cell Physiol , vol.52 , pp. 1856-1871
    • Zhong, R.1    Lee, C.2    McCarthy, R.L.3    Reeves, C.K.4    Jones, E.G.5
  • 20
    • 84875192888 scopus 로고    scopus 로고
    • Regulation of secondary wall synthesis and cell death by NAC transcription factors in the monocot Brachypodium distachyon
    • Valdivia ER, Herrera MT, Gianzo C, Fidalgo J, Revilla G, et al. (2013) Regulation of secondary wall synthesis and cell death by NAC transcription factors in the monocot Brachypodium distachyon. J Exp Bot 64: 1333-1343.
    • (2013) J Exp Bot , vol.64 , pp. 1333-1343
    • Valdivia, E.R.1    Herrera, M.T.2    Gianzo, C.3    Fidalgo, J.4    Revilla, G.5
  • 21
    • 84875186825 scopus 로고    scopus 로고
    • Transcriptional regulation of grass secondary cell wall biosynthesis: playing catch-up with Arabidopsis thaliana. Front Plant Sci
    • Handakumbura PP, Hazen SP, (2012) Transcriptional regulation of grass secondary cell wall biosynthesis: playing catch-up with Arabidopsis thaliana. Front Plant Sci. 3: 74.
    • (2012) , vol.3 , pp. 74
    • Handakumbura, P.P.1    Hazen, S.P.2
  • 22
    • 27144549989 scopus 로고    scopus 로고
    • EgMYB2, a new transcriptional activator from Eucalyptus xylem, regulates secondary cell wall formation and lignin biosynthesis
    • Goicoechea M, Lacombe E, Legay S, Mihaljevic S, Rech P, et al. (2005) EgMYB2, a new transcriptional activator from Eucalyptus xylem, regulates secondary cell wall formation and lignin biosynthesis. Plant J 43: 553-567.
    • (2005) Plant J , vol.43 , pp. 553-567
    • Goicoechea, M.1    Lacombe, E.2    Legay, S.3    Mihaljevic, S.4    Rech, P.5
  • 23
  • 24
    • 77953557291 scopus 로고    scopus 로고
    • The poplar MYB transcription factors, PtrMYB3 and PtrMYB20, are involved in the regulation of secondary wall biosynthesis
    • McCarthy RL, Zhong R, Fowler S, Lyskowski D, Piyasena H, et al. (2010) The poplar MYB transcription factors, PtrMYB3 and PtrMYB20, are involved in the regulation of secondary wall biosynthesis. Plant Cell Physiol 51: 1084-1090.
    • (2010) Plant Cell Physiol , vol.51 , pp. 1084-1090
    • McCarthy, R.L.1    Zhong, R.2    Fowler, S.3    Lyskowski, D.4    Piyasena, H.5
  • 25
    • 84857226184 scopus 로고    scopus 로고
    • MYB46 and MYB83 bind to the SMRE sites and directly activate a suite of transcription factors and secondary wall biosynthetic genes
    • Zhong R, Ye Z-H, (2012) MYB46 and MYB83 bind to the SMRE sites and directly activate a suite of transcription factors and secondary wall biosynthetic genes. Plant Cell Physiol 53: 368-380.
    • (2012) Plant Cell Physiol , vol.53 , pp. 368-380
    • Zhong, R.1    Ye, Z.-H.2
  • 26
    • 84856880107 scopus 로고    scopus 로고
    • Identification of a cis-acting regulatory motif recognized by MYB46, a master transcriptional regulator of secondary wall biosynthesis
    • Kim WC, Ko JH, Han KH, (2012) Identification of a cis-acting regulatory motif recognized by MYB46, a master transcriptional regulator of secondary wall biosynthesis. Plant Mol Biol 78: 489-501.
    • (2012) Plant Mol Biol , vol.78 , pp. 489-501
    • Kim, W.C.1    Ko, J.H.2    Han, K.H.3
  • 27
    • 0142035247 scopus 로고    scopus 로고
    • Genome-wide characterization of the lignification toolbox in Arabidopsis
    • Raes J, Rohde A, Christensen JH, Peer YV, Boerjan W, (2003) Genome-wide characterization of the lignification toolbox in Arabidopsis. Plant Physiol 133: 1051-1071.
    • (2003) Plant Physiol , vol.133 , pp. 1051-1071
    • Raes, J.1    Rohde, A.2    Christensen, J.H.3    Peer, Y.V.4    Boerjan, W.5
  • 28
    • 4644291602 scopus 로고    scopus 로고
    • The genetic control of lignin deposition during plant growth and development
    • Rogers LA, Campbell MM, (2004) The genetic control of lignin deposition during plant growth and development. New Phytol 164: 17-30.
    • (2004) New Phytol , vol.164 , pp. 17-30
    • Rogers, L.A.1    Campbell, M.M.2
  • 29
    • 70450230537 scopus 로고    scopus 로고
    • MYB83 is a direct target of SND1 and acts redundantly with MYB46 in the regulation of secondary cell wall biosynthesis in Arabidopsis
    • McCarthy RL, Zhong R, Ye Z-H, (2009) MYB83 is a direct target of SND1 and acts redundantly with MYB46 in the regulation of secondary cell wall biosynthesis in Arabidopsis. Plant Cell Physiol 50: 1950-1964.
    • (2009) Plant Cell Physiol , vol.50 , pp. 1950-1964
    • McCarthy, R.L.1    Zhong, R.2    Ye, Z.-H.3
  • 30
    • 35748936175 scopus 로고    scopus 로고
    • The MYB46 transcription factor is a direct target of SND1 and regulates secondary wall biosynthesis in Arabidopsis
    • Zhong R, Richardson EA, Ye Z-H, (2007) The MYB46 transcription factor is a direct target of SND1 and regulates secondary wall biosynthesis in Arabidopsis. Plant Cell 19: 2776-2792.
    • (2007) Plant Cell , vol.19 , pp. 2776-2792
    • Zhong, R.1    Richardson, E.A.2    Ye, Z.-H.3
  • 31
    • 3543007738 scopus 로고    scopus 로고
    • Identification of the minimal repression domain of SUPERMAN shows that the DLELRL hexapeptide is both necessary and sufficient for repression of transcription in Arabidopsis
    • Hiratsu K, Mitsuda N, Matsui K, Ohme-Takagi M, (2004) Identification of the minimal repression domain of SUPERMAN shows that the DLELRL hexapeptide is both necessary and sufficient for repression of transcription in Arabidopsis. Biochem Biophy Res Commun 321: 172-178.
    • (2004) Biochem Biophy Res Commun , vol.321 , pp. 172-178
    • Hiratsu, K.1    Mitsuda, N.2    Matsui, K.3    Ohme-Takagi, M.4
  • 32
    • 80051753694 scopus 로고    scopus 로고
    • Molecular dissection of xylan biosynthesis during wood formation in poplar
    • Lee C, Teng Q, Zhong R, Ye Z-H, (2011) Molecular dissection of xylan biosynthesis during wood formation in poplar. Mol Plant 4: 730-747.
    • (2011) Mol Plant , vol.4 , pp. 730-747
    • Lee, C.1    Teng, Q.2    Zhong, R.3    Ye, Z.-H.4
  • 33
    • 78649862688 scopus 로고    scopus 로고
    • Global analysis of direct targets of secondary wall NAC master switches in Arabidopsis
    • Zhong R, Lee C, Ye Z-H, (2010) Global analysis of direct targets of secondary wall NAC master switches in Arabidopsis. Mol Plant 3: 1087-1103.
    • (2010) Mol Plant , vol.3 , pp. 1087-1103
    • Zhong, R.1    Lee, C.2    Ye, Z.-H.3
  • 34
    • 62549142281 scopus 로고    scopus 로고
    • MYB58 and MYB63 are transcriptional activators of the lignin biosynthetic pathway during secondary cell wall formation in Arabidopsis
    • Zhou J, Lee C, Zhong R, Ye Z-H, (2009) MYB58 and MYB63 are transcriptional activators of the lignin biosynthetic pathway during secondary cell wall formation in Arabidopsis. Plant Cell 21: 248-266.
    • (2009) Plant Cell , vol.21 , pp. 248-266
    • Zhou, J.1    Lee, C.2    Zhong, R.3    Ye, Z.-H.4
  • 35
    • 74549176584 scopus 로고    scopus 로고
    • Towards a systems approach for lignin biosynthesis in Populus trichocarpa: transcript abundance and specificity of the monolignol biosynthetic genes
    • Shi R, Sun YH, Li Q, Heber S, Sederoff R, et al. (2010) Towards a systems approach for lignin biosynthesis in Populus trichocarpa: transcript abundance and specificity of the monolignol biosynthetic genes. Plant Cell Physiol. 51, 144-163.
    • (2010) Plant Cell Physiol , vol.51 , pp. 144-163
    • Shi, R.1    Sun, Y.H.2    Li, Q.3    Heber, S.4    Sederoff, R.5
  • 36
    • 55249088062 scopus 로고    scopus 로고
    • Involvement of Pinus taeda MYB1 and MYB8 in phenylpropanoid metabolism and secondary cell wall biogenesis: a comparative in planta analysis
    • Bomal C, Bedon F, Caron S, Mansfield SD, Levasseur C, et al. (2008) Involvement of Pinus taeda MYB1 and MYB8 in phenylpropanoid metabolism and secondary cell wall biogenesis: a comparative in planta analysis. J Exp Bot 59: 3925-3939.
    • (2008) J Exp Bot , vol.59 , pp. 3925-3939
    • Bomal, C.1    Bedon, F.2    Caron, S.3    Mansfield, S.D.4    Levasseur, C.5
  • 37
    • 67349284106 scopus 로고    scopus 로고
    • The maize ZmMYB42 represses the phenylpropanoid pathway and affects the cell wall structure, composition and degradability in Arabidopsis thaliana
    • Sonbol FM, Fornalé S, Capellades M, Encina A, Touriño S, et al. (2009) The maize ZmMYB42 represses the phenylpropanoid pathway and affects the cell wall structure, composition and degradability in Arabidopsis thaliana. Plant Mol Biol 70: 283-296.
    • (2009) Plant Mol Biol , vol.70 , pp. 283-296
    • Sonbol, F.M.1    Fornalé, S.2    Capellades, M.3    Encina, A.4    Touriño, S.5
  • 38
    • 77954058215 scopus 로고    scopus 로고
    • Transcriptional regulation of lignin biosynthesis
    • Zhong R, Ye Z-H, (2009) Transcriptional regulation of lignin biosynthesis. Plant Signal Behav 4: 1028-1034.
    • (2009) Plant Signal Behav , vol.4 , pp. 1028-1034
    • Zhong, R.1    Ye, Z.-H.2
  • 39
    • 78449255457 scopus 로고    scopus 로고
    • ZmMYB31 directly represses maize lignin genes and redirects the phenylpropanoid metabolic flux
    • Fornalé S, Shi X, Chai C, Encina A, Irar S, et al. (2010) ZmMYB31 directly represses maize lignin genes and redirects the phenylpropanoid metabolic flux. Plant J 64: 633-644.
    • (2010) Plant J , vol.64 , pp. 633-644
    • Fornalé, S.1    Shi, X.2    Chai, C.3    Encina, A.4    Irar, S.5
  • 40
    • 34748821912 scopus 로고    scopus 로고
    • Molecular characterization of EgMYB1, a putative transcriptional repressor of the lignin biosynthetic pathway
    • Legay S, Lacombe E, Goicoechea M, Briere C, Seguin A, et al. (2007) Molecular characterization of EgMYB1, a putative transcriptional repressor of the lignin biosynthetic pathway. Plant Sci 173: 542-549.
    • (2007) Plant Sci , vol.173 , pp. 542-549
    • Legay, S.1    Lacombe, E.2    Goicoechea, M.3    Briere, C.4    Seguin, A.5
  • 42
    • 0000122573 scopus 로고
    • PHYLIP - Phylogeny Inference Package (Version 3.2)
    • Felsenstein J, (1989) PHYLIP - Phylogeny Inference Package (Version 3.2). Cladistics 5: 164-166.
    • (1989) Cladistics , vol.5 , pp. 164-166
    • Felsenstein, J.1
  • 43
    • 0030203863 scopus 로고    scopus 로고
    • TREEVIEW: An application to display phylogenetic trees on personal computers
    • Page RDM, (1996) TREEVIEW: An application to display phylogenetic trees on personal computers. Computer Appl Biosci 12: 357-358.
    • (1996) Computer Appl Biosci , vol.12 , pp. 357-358
    • Page, R.D.M.1
  • 45
    • 33646690468 scopus 로고    scopus 로고
    • Disruption of cortical microtubules by overexpression of green fluorescent protein-tagged α-tubulin 6 causes a marked reduction in cell wall synthesis
    • Burk DH, Zhong R, Morrison WH III, Ye Z-H, (2006) Disruption of cortical microtubules by overexpression of green fluorescent protein-tagged α-tubulin 6 causes a marked reduction in cell wall synthesis. J Integr Plant Biol 48: 85-98.
    • (2006) J Integr Plant Biol , vol.48 , pp. 85-98
    • Burk, D.H.1    Zhong, R.2    Morrison III, W.H.3    Ye, Z.-H.4
  • 46
    • 33845736876 scopus 로고    scopus 로고
    • SND1, a NAC domain transcription factor, is a key regulator of secondary wall synthesis in fibers of Arabidopsis
    • Zhong R, Demura T, Ye Z-H, (2006) SND1, a NAC domain transcription factor, is a key regulator of secondary wall synthesis in fibers of Arabidopsis. Plant Cell 18: 3158-3170.
    • (2006) Plant Cell , vol.18 , pp. 3158-3170
    • Zhong, R.1    Demura, T.2    Ye, Z.-H.3
  • 47
    • 84907123778 scopus 로고
    • The use of an optical brightener in the study of plant structure
    • Hughes J, McCully ME, (1975) The use of an optical brightener in the study of plant structure. Stain Technol 50: 319-329.
    • (1975) Stain Technol , vol.50 , pp. 319-329
    • Hughes, J.1    McCully, M.E.2
  • 48
    • 16244383185 scopus 로고    scopus 로고
    • Monoclonal antibodies to plant cell wall xylans and arabinoxylans
    • McCartney L, Marcus SE, Knox JP, (2005) Monoclonal antibodies to plant cell wall xylans and arabinoxylans. J Histochem Cytochem 53: 543-546.
    • (2005) J Histochem Cytochem , vol.53 , pp. 543-546
    • McCartney, L.1    Marcus, S.E.2    Knox, J.P.3
  • 49
    • 85047682075 scopus 로고    scopus 로고
    • Signal transduction in maize and Arabidopsis mesophyll protoplasts
    • Sheen J, (2001) Signal transduction in maize and Arabidopsis mesophyll protoplasts. Plant Physiol 127: 1466-1475.
    • (2001) Plant Physiol , vol.127 , pp. 1466-1475
    • Sheen, J.1
  • 50
    • 0035971183 scopus 로고    scopus 로고
    • Functional interactions of lanthanum and phospholipase D with the abscisic acid signaling effectors VP1 and ABI1-1 in rice protoplasts
    • Gampala SS, Hagenbeek D, Rock CD, (2001) Functional interactions of lanthanum and phospholipase D with the abscisic acid signaling effectors VP1 and ABI1-1 in rice protoplasts. J Biol Chem 276: 9855-9860.
    • (2001) J Biol Chem , vol.276 , pp. 9855-9860
    • Gampala, S.S.1    Hagenbeek, D.2    Rock, C.D.3


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