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Volumn 195, Issue 12, 2013, Pages 2864-2879

Evidence that a metabolic microcompartment contains and recycles private cofactor pools

Author keywords

[No Author keywords available]

Indexed keywords

ACETALDEHYDE; ACETYL COENZYME A; ACETYL PHOSPHATE; ALCOHOL; ASTRINGENT AGENT; BACTERIAL PROTEIN; COBAMAMIDE; COENZYME A; CYANOCOBALAMIN; ETHANOLAMINE; EUTD PROTEIN; EUTE PROTEIN; EUTG PROTEIN; MUTANT PROTEIN; NICOTINAMIDE ADENINE DINUCLEOTIDE; UNCLASSIFIED DRUG; VOLATILE AGENT;

EID: 84880017088     PISSN: 00219193     EISSN: 10985530     Source Type: Journal    
DOI: 10.1128/JB.02179-12     Document Type: Article
Times cited : (56)

References (56)
  • 2
    • 0032851358 scopus 로고    scopus 로고
    • The 17-gene ethanolamine (eut) operon of Salmonella typhimurium encodes five homologues of carboxysome shell proteins
    • Kofoid E, Rappleye C, Stojiljkovic I, Roth J. 1999. The 17-gene ethanolamine (eut) operon of Salmonella typhimurium encodes five homologues of carboxysome shell proteins. J. Bacteriol. 181:5317-5329.
    • (1999) J. Bacteriol. , vol.181 , pp. 5317-5329
    • Kofoid, E.1    Rappleye, C.2    Stojiljkovic, I.3    Roth, J.4
  • 3
    • 0015816557 scopus 로고
    • Functional organelles in prokaryotes: polyhedral inclusions (carboxysomes) of Thiobacillus neapolitanus
    • Shively JM, Ball F, Brown DH, Saunders RE. 1973. Functional organelles in prokaryotes: polyhedral inclusions (carboxysomes) of Thiobacillus neapolitanus. Science 182:584-586.
    • (1973) Science , vol.182 , pp. 584-586
    • Shively, J.M.1    Ball, F.2    Brown, D.H.3    Saunders, R.E.4
  • 4
    • 33645964606 scopus 로고    scopus 로고
    • Conserving a volatile metabolite: a role for carboxysome-like organelles in Salmonella enterica
    • Penrod JT, Roth JR. 2006. Conserving a volatile metabolite: a role for carboxysome-like organelles in Salmonella enterica. J. Bacteriol. 188: 2865-2874.
    • (2006) J. Bacteriol. , vol.188 , pp. 2865-2874
    • Penrod, J.T.1    Roth, J.R.2
  • 5
    • 41949137665 scopus 로고    scopus 로고
    • 12-dependent 1,2-propanediol degradation provide protection from DNA and cellular damage by a reactive metabolic intermediate
    • 12-dependent 1,2-propanediol degradation provide protection from DNA and cellular damage by a reactive metabolic intermediate. J. Bacteriol. 190:2966-2971.
    • (2008) J. Bacteriol. , vol.190 , pp. 2966-2971
    • Sampson, E.M.1    Bobik, T.A.2
  • 6
    • 0024075956 scopus 로고
    • Ethanolamine utilization in Salmonella typhimurium
    • Roof DM, Roth JR. 1988. Ethanolamine utilization in Salmonella typhimurium. J. Bacteriol. 170:3855-3863.
    • (1988) J. Bacteriol. , vol.170 , pp. 3855-3863
    • Roof, D.M.1    Roth, J.R.2
  • 7
    • 0024393078 scopus 로고
    • 12-dependent ethanolamine utilization in Salmonella typhimurium
    • 12-dependent ethanolamine utilization in Salmonella typhimurium. J. Bacteriol. 171: 3316-3323.
    • (1989) J. Bacteriol. , vol.171 , pp. 3316-3323
    • Roof, D.M.1    Roth, J.R.2
  • 8
    • 80052818704 scopus 로고    scopus 로고
    • Coproduction of acetaldehyde and hydrogen during glucose fermentation by Escherichia coli
    • Zhu H, Gonzalez R, Bobik TA. 2011. Coproduction of acetaldehyde and hydrogen during glucose fermentation by Escherichia coli. Appl. Environ. Microbiol. 77:6441-6450.
    • (2011) Appl. Environ. Microbiol. , vol.77 , pp. 6441-6450
    • Zhu, H.1    Gonzalez, R.2    Bobik, T.A.3
  • 9
    • 1542286908 scopus 로고    scopus 로고
    • The eutD gene of Salmonella enterica encodes a protein with phosphotransacetylase enzyme activity
    • Brinsmade SR, Escalante-Semerena JC. 2004. The eutD gene of Salmonella enterica encodes a protein with phosphotransacetylase enzyme activity. J. Bacteriol. 186:1890-1892.
    • (2004) J. Bacteriol. , vol.186 , pp. 1890-1892
    • Brinsmade, S.R.1    Escalante-Semerena, J.C.2
  • 10
    • 79951518426 scopus 로고    scopus 로고
    • Characterization of Escherichia coli EutD: a phosphotransacetylase of the ethanolamine operon
    • Bologna F, Campos-Bermudez V, Saavedra D, Andreo C, Drincovich M. 2010. Characterization of Escherichia coli EutD: a phosphotransacetylase of the ethanolamine operon. J. Microbiol. 48:629-636.
    • (2010) J. Microbiol. , vol.48 , pp. 629-636
    • Bologna, F.1    Campos-Bermudez, V.2    Saavedra, D.3    Andreo, C.4    Drincovich, M.5
  • 11
    • 0035094993 scopus 로고    scopus 로고
    • 12-dependent anaerobic growth of Salmonella enterica serovar Typhimurium on ethanolamine or 1,2-propanediol
    • 12-dependent anaerobic growth of Salmonella enterica serovar Typhimurium on ethanolamine or 1,2-propanediol. J. Bacteriol. 183:2463-2475.
    • (2001) J. Bacteriol. , vol.183 , pp. 2463-2475
    • Price-Carter, M.1    Tingey, J.2    Bobik, T.A.3    Roth, J.R.4
  • 12
    • 27744588276 scopus 로고    scopus 로고
    • Acetate excretion during growth of Salmonella enterica on ethanolamine requires phosphotransacetylase (EutD) activity, and acetate recapture requires acetyl-CoA synthetase (Acs) and phosphotransacetylase (Pta) activities
    • Starai VJ, Garrity J, Escalante-Semerena JC. 2005. Acetate excretion during growth of Salmonella enterica on ethanolamine requires phosphotransacetylase (EutD) activity, and acetate recapture requires acetyl-CoA synthetase (Acs) and phosphotransacetylase (Pta) activities. Microbiology 151:3793-3801.
    • (2005) Microbiology , vol.151 , pp. 3793-3801
    • Starai, V.J.1    Garrity, J.2    Escalante-Semerena, J.C.3
  • 13
    • 28044459168 scopus 로고    scopus 로고
    • Minimal functions and physiological conditions required for growth of Salmonella enterica on ethanolamine in the absence of the metabolosome
    • Brinsmade SR, Paldon T, Escalante-Semerena JC. 2005. Minimal functions and physiological conditions required for growth of Salmonella enterica on ethanolamine in the absence of the metabolosome. J. Bacteriol. 187:8039-8046.
    • (2005) J. Bacteriol. , vol.187 , pp. 8039-8046
    • Brinsmade, S.R.1    Paldon, T.2    Escalante-Semerena, J.C.3
  • 14
    • 80053580487 scopus 로고    scopus 로고
    • The N-terminal region of the medium subunit (PduD) packages adenosylcobalamin-dependent diol dehydratase (PduCDE) into the Pdu microcompartment
    • Fan C, Bobik TA. 2011. The N-terminal region of the medium subunit (PduD) packages adenosylcobalamin-dependent diol dehydratase (PduCDE) into the Pdu microcompartment. J. Bacteriol. 193:5623-5628.
    • (2011) J. Bacteriol. , vol.193 , pp. 5623-5628
    • Fan, C.1    Bobik, T.A.2
  • 15
    • 84866280237 scopus 로고    scopus 로고
    • Interactions between the termini of lumen enzymes and shell proteins mediate enzyme encapsulation into bacterial microcompartments
    • Fan C, Cheng S, Sinha S, Bobik TA. 2012. Interactions between the termini of lumen enzymes and shell proteins mediate enzyme encapsulation into bacterial microcompartments. Proc. Natl. Acad. Sci. U. S. A. 109:14995-15000.
    • (2012) Proc. Natl. Acad. Sci. U. S. A. , vol.109 , pp. 14995-15000
    • Fan, C.1    Cheng, S.2    Sinha, S.3    Bobik, T.A.4
  • 17
    • 0003455675 scopus 로고
    • Advanced bacterial genetics
    • Cold Spring Harbor Laboratory, Cold Spring Harbor, NY
    • Davis RW, Botstein D, Roth JR. 1980. Advanced bacterial genetics. Cold Spring Harbor Laboratory, Cold Spring Harbor, NY.
    • (1980)
    • Davis, R.W.1    Botstein, D.2    Roth, J.R.3
  • 18
    • 0014992845 scopus 로고
    • A method for detection of phage mutants with altered transducing ability
    • Schmieger H. 1971. A method for detection of phage mutants with altered transducing ability. Mol. Gen. Genet. 110:378-381.
    • (1971) Mol. Gen. Genet. , vol.110 , pp. 378-381
    • Schmieger, H.1
  • 19
    • 4944235152 scopus 로고    scopus 로고
    • A pH-sensitive function and phenotype: evidence that EutH facilitates diffusion of uncharged ethanolamine in Salmonella enterica
    • Penrod JT, Mace CC, Roth JR. 2004. A pH-sensitive function and phenotype: evidence that EutH facilitates diffusion of uncharged ethanolamine in Salmonella enterica. J. Bacteriol. 186:6885-6890.
    • (2004) J. Bacteriol. , vol.186 , pp. 6885-6890
    • Penrod, J.T.1    Mace, C.C.2    Roth, J.R.3
  • 20
    • 0034612342 scopus 로고    scopus 로고
    • One-step inactivation of chromosomal genes in Escherichia coli K-12 using PCR products
    • Datsenko KA, Wanner BL. 2000. One-step inactivation of chromosomal genes in Escherichia coli K-12 using PCR products. Proc. Natl. Acad. Sci. U. S. A. 97:6640-6645.
    • (2000) Proc. Natl. Acad. Sci. U. S. A. , vol.97 , pp. 6640-6645
    • Datsenko, K.A.1    Wanner, B.L.2
  • 21
    • 0038472057 scopus 로고    scopus 로고
    • Recombineering with overlapping single-stranded DNA oligonucleotides: testing a recombination intermediate
    • Yu D, Sawitzke JA, Ellis H, Court DL. 2003. Recombineering with overlapping single-stranded DNA oligonucleotides: testing a recombination intermediate. Proc. Natl. Acad. Sci. U. S. A. 100:7207-7212.
    • (2003) Proc. Natl. Acad. Sci. U. S. A. , vol.100 , pp. 7207-7212
    • Yu, D.1    Sawitzke, J.A.2    Ellis, H.3    Court, D.L.4
  • 22
    • 0028944587 scopus 로고
    • MudSacI, a transposon with strong selectable and counterselectable markers: use for rapid mapping of chromosomal mutations in Salmonella typhimurium
    • Lawes M, Maloy S. 1995. MudSacI, a transposon with strong selectable and counterselectable markers: use for rapid mapping of chromosomal mutations in Salmonella typhimurium. J. Bacteriol. 177:1383-1387.
    • (1995) J. Bacteriol. , vol.177 , pp. 1383-1387
    • Lawes, M.1    Maloy, S.2
  • 25
    • 0026657140 scopus 로고
    • Autogenous regulation of ethanolamine utilization by a transcriptional activator of the eut operon in Salmonella typhimurium
    • Roof DM, Roth JR. 1992. Autogenous regulation of ethanolamine utilization by a transcriptional activator of the eut operon in Salmonella typhimurium. J. Bacteriol. 174:6634-6643.
    • (1992) J. Bacteriol. , vol.174 , pp. 6634-6643
    • Roof, D.M.1    Roth, J.R.2
  • 26
    • 0003785155 scopus 로고
    • Experiments in molecular genetics
    • Cold Spring Harbor Laboratory, Cold Spring Harbor, NY.
    • Miller JH. 1972. Experiments in molecular genetics. Cold Spring Harbor Laboratory, Cold Spring Harbor, NY.
    • (1972)
    • Miller, J.H.1
  • 27
    • 0028262935 scopus 로고
    • A rationale for autoinduction of a transcriptional activator: ethanolamine ammonia-lyase (EutBC) and the operon activator (EutR) compete for adenosyl-cobalamin in Salmonella typhimurium
    • Sheppard DE, Roth JR. 1994. A rationale for autoinduction of a transcriptional activator: ethanolamine ammonia-lyase (EutBC) and the operon activator (EutR) compete for adenosyl-cobalamin in Salmonella typhimurium. J. Bacteriol. 176:1287-1296.
    • (1994) J. Bacteriol. , vol.176 , pp. 1287-1296
    • Sheppard, D.E.1    Roth, J.R.2
  • 28
    • 0019996140 scopus 로고
    • Model for the participation of quasi-palindromic DNA sequences in frameshift mutation
    • Ripley LS. 1982. Model for the participation of quasi-palindromic DNA sequences in frameshift mutation. Proc. Natl. Acad. Sci. U. S. A. 79:4128-4132.
    • (1982) Proc. Natl. Acad. Sci. U. S. A. , vol.79 , pp. 4128-4132
    • Ripley, L.S.1
  • 29
    • 0021243178 scopus 로고
    • Conditionally transposition-defective derivative of Mu d1(Amp Lac)
    • Hughes KT, Roth JR. 1984. Conditionally transposition-defective derivative of Mu d1(Amp Lac). J. Bacteriol. 159:130-137.
    • (1984) J. Bacteriol. , vol.159 , pp. 130-137
    • Hughes, K.T.1    Roth, J.R.2
  • 31
    • 0035838544 scopus 로고    scopus 로고
    • Novel alcohol dehydrogenase activity in a mutant of Salmonella able to use ethanol as sole carbon source
    • Dailly Y, Mat-Jan F, Clark DP. 2001. Novel alcohol dehydrogenase activity in a mutant of Salmonella able to use ethanol as sole carbon source. FEMS Microbiol. Lett. 201:41-45.
    • (2001) FEMS Microbiol. Lett , vol.201 , pp. 41-45
    • Dailly, Y.1    Mat-Jan, F.2    Clark, D.P.3
  • 32
    • 0033641955 scopus 로고    scopus 로고
    • Comparison of the fermentative alcohol dehydrogenases of Salmonella typhimurium and Escherichia coli
    • Dailly YP, Bunch P, Clark DP. 2000. Comparison of the fermentative alcohol dehydrogenases of Salmonella typhimurium and Escherichia coli. Microbios 103:179-196.
    • (2000) Microbios , vol.103 , pp. 179-196
    • Dailly, Y.P.1    Bunch, P.2    Clark, D.P.3
  • 35
    • 0017759929 scopus 로고
    • The enzymic interconversion of acetate and acetyl-coenzyme A in Escherichia coli
    • Brown TDK, Jones-Mortimer MC, Kornberg HL. 1977. The enzymic interconversion of acetate and acetyl-coenzyme A in Escherichia coli. J. Gen. Microbiol. 102:327-336.
    • (1977) J. Gen. Microbiol. , vol.102 , pp. 327-336
    • Brown, T.D.K.1    Jones-Mortimer, M.C.2    Kornberg, H.L.3
  • 36
    • 0018950458 scopus 로고
    • Isolation and characterization of acetate kinase and phosphotransacetylase mutants of Escherichia coli and Salmonella typhimurium
    • LeVine SM, Ardeshir F, Ames GF. 1980. Isolation and characterization of acetate kinase and phosphotransacetylase mutants of Escherichia coli and Salmonella typhimurium. J. Bacteriol. 143:1081-1085.
    • (1980) J. Bacteriol. , vol.143 , pp. 1081-1085
    • LeVine, S.M.1    Ardeshir, F.2    Ames, G.F.3
  • 37
    • 0000146513 scopus 로고    scopus 로고
    • Two-carbon compounds and fatty acids as carbon sources
    • In Neidhardt FC, Curtiss R, III, Ingraham JL, Lin ECC, Low KB, Magasanik B, Reznikoff WS, Riley M, Schaechter M, Umbarger HE. (ed), 2nd ed, vol 1 American Society for Microbiology, Washington, DC.
    • Clark DP, Cronan JP, Jr. 1996. Two-carbon compounds and fatty acids as carbon sources, p 343-357. In Neidhardt FC, Curtiss R, III, Ingraham JL, Lin ECC, Low KB, Magasanik B, Reznikoff WS, Riley M, Schaechter M, Umbarger HE. (ed), Escherichia coli and Salmonella: cellular and molecular biology, 2nd ed, vol 1. American Society for Microbiology, Washington, DC.
    • (1996) Escherichia coli and Salmonella. cellular and molecular biology , pp. 343-357
    • Clark, D.P.1    Cronan Jr., J.P.2
  • 38
    • 0029009026 scopus 로고
    • Cloning, characterization, and functional expression of acs, the gene which encodes acetyl coenzyme A synthetase in Escherichia coli
    • Kumari S, Tishel R, Eisenbach M, Wolfe AJ. 1995. Cloning, characterization, and functional expression of acs, the gene which encodes acetyl coenzyme A synthetase in Escherichia coli. J. Bacteriol. 177:2878-2886.
    • (1995) J. Bacteriol. , vol.177 , pp. 2878-2886
    • Kumari, S.1    Tishel, R.2    Eisenbach, M.3    Wolfe, A.J.4
  • 39
    • 0037560995 scopus 로고    scopus 로고
    • 12-dependent isomerization (eliminating) reactions
    • 12-dependent isomerization (eliminating) reactions. Chem. Rev. 103:2095-2128.
    • (2003) Chem. Rev. , vol.103 , pp. 2095-2128
    • Toraya, T.1
  • 40
    • 4944253783 scopus 로고    scopus 로고
    • Identification of a reactivating factor for adenosylcobalamin-dependent ethanolamine ammonia lyase
    • Mori K, Bando R, Hieda N, Toraya T. 2004. Identification of a reactivating factor for adenosylcobalamin-dependent ethanolamine ammonia lyase. J. Bacteriol. 186:6845-6854.
    • (2004) J. Bacteriol. , vol.186 , pp. 6845-6854
    • Mori, K.1    Bando, R.2    Hieda, N.3    Toraya, T.4
  • 41
    • 0018850564 scopus 로고
    • Escherichia coli mutants with altered control of alcohol dehydrogenase and nitrate reductase
    • Clark D, Cronan JE. 1980. Escherichia coli mutants with altered control of alcohol dehydrogenase and nitrate reductase. J. Bacteriol. 141:177-183.
    • (1980) J. Bacteriol. , vol.141 , pp. 177-183
    • Clark, D.1    Cronan, J.E.2
  • 44
    • 0029671428 scopus 로고    scopus 로고
    • 12 synthesis among enteric bacteria: evidence for loss and reacquisition of a multigene complex
    • 12 synthesis among enteric bacteria: evidence for loss and reacquisition of a multigene complex. Genetics 142:11-24.
    • (1996) Genetics , vol.142 , pp. 11-24
    • Lawrence, J.G.1    Roth, J.R.2
  • 45
    • 28444474563 scopus 로고    scopus 로고
    • Identification of core and variable components of the Salmonella enterica subspecies I genome by microarray
    • Anjum MF, Marooney C, Fookes M, Baker S, Dougan G, Ivens A, Woodward MJ. 2005. Identification of core and variable components of the Salmonella enterica subspecies I genome by microarray. Infect. Immun. 73:7894-7905.
    • (2005) Infect. Immun. , vol.73 , pp. 7894-7905
    • Anjum, M.F.1    Marooney, C.2    Fookes, M.3    Baker, S.4    Dougan, G.5    Ivens, A.6    Woodward, M.J.7
  • 46
    • 0022745252 scopus 로고
    • Growth of salmonellas in different enrichment media
    • Patil MD, Parhad NM. 1986. Growth of salmonellas in different enrichment media. J. Appl. Microbiol. 61:19-24.
    • (1986) J. Appl. Microbiol. , vol.61 , pp. 19-24
    • Patil, M.D.1    Parhad, N.M.2
  • 47
    • 0025141627 scopus 로고
    • New plate medium for facilitated differentiation of Salmonella spp. from Proteus spp. and other enteric bacteria.
    • Rambach A. 1990. New plate medium for facilitated differentiation of Salmonella spp. from Proteus spp. and other enteric bacteria. Appl. Environ. Microbiol. 56:301-303.
    • (1990) Appl. Environ. Microbiol. , vol.56 , pp. 301-303
    • Rambach, A.1
  • 48
    • 4344682568 scopus 로고    scopus 로고
    • The eutT gene of Salmonella enterica encodes an oxygen-labile, metal-containing ATP: corrinoid adenosyltransferase enzyme
    • Buan NR, Suh S-J, Escalante-Semerena JC. 2004. The eutT gene of Salmonella enterica encodes an oxygen-labile, metal-containing ATP: corrinoid adenosyltransferase enzyme. J. Bacteriol. 186:5708-5714.
    • (2004) J. Bacteriol. , vol.186 , pp. 5708-5714
    • Buan, N.R.1    Suh, S.-J.2    Escalante-Semerena, J.C.3
  • 50
    • 0042389658 scopus 로고    scopus 로고
    • 12-dependent degradation of 1,2-propanediol in Salmonella enterica serovar Typhimurium LT2
    • 12-dependent degradation of 1,2-propanediol in Salmonella enterica serovar Typhimurium LT2. J. Bacteriol. 185:5086-5095.
    • (2003) J. Bacteriol. , vol.185 , pp. 5086-5095
    • Havemann, G.D.1    Bobik, T.A.2
  • 51
    • 77957353322 scopus 로고    scopus 로고
    • Characterization of the PduS cobalamin reductase of Salmonella enterica and its role in the Pdu microcompartment
    • Cheng S, Bobik TA. 2010. Characterization of the PduS cobalamin reductase of Salmonella enterica and its role in the Pdu microcompartment. J. Bacteriol. 192:5071-5080.
    • (2010) J. Bacteriol. , vol.192 , pp. 5071-5080
    • Cheng, S.1    Bobik, T.A.2
  • 52
    • 74849140487 scopus 로고    scopus 로고
    • Structure and mechanisms of a protein-based organelle in Escherichia coli
    • Tanaka S, Sawaya MR, Yeates TO. 2010. Structure and mechanisms of a protein-based organelle in Escherichia coli. Science 327:81-84.
    • (2010) Science , vol.327 , pp. 81-84
    • Tanaka, S.1    Sawaya, M.R.2    Yeates, T.O.3
  • 54
    • 78549246253 scopus 로고    scopus 로고
    • Structural insight into the mechanisms of transport across the Salmonella enterica Pdu microcompartment shell
    • Crowley CS, Cascio D, Sawaya MR, Kopstein JS, Bobik TA, Yeates TO. 2010. Structural insight into the mechanisms of transport across the Salmonella enterica Pdu microcompartment shell. J. Biol. Chem. 285:37838-37846.
    • (2010) J. Biol. Chem. , vol.285 , pp. 37838-37846
    • Crowley, C.S.1    Cascio, D.2    Sawaya, M.R.3    Kopstein, J.S.4    Bobik, T.A.5    Yeates, T.O.6
  • 55
    • 55849129963 scopus 로고    scopus 로고
    • Halothiobacillus neapolitanus carboxysomes sequester heterologous and chimeric RuBisCo species
    • doi:10.1371/journal.pone .0003570
    • Menon BB, Dou Z, Heinhorst S, Shively JM, Cannon GC. 2008. Halothiobacillus neapolitanus carboxysomes sequester heterologous and chimeric RuBisCo species. PLoS One 3:e3570. doi:10.1371/journal.pone .0003570.
    • (2008) PLoS One , vol.3
    • Menon, B.B.1    Dou, Z.2    Heinhorst, S.3    Shively, J.M.4    Cannon, G.C.5
  • 56
    • 84867515386 scopus 로고    scopus 로고
    • The PduQ enzyme is an alcohol dehydrogenase used to recycle NAD internally within the Pdu microcompartment of Salmonella enterica
    • doi:10 .1371/journal.pone.0047144
    • Cheng S, Fand C, Sinha S, Bobik TA. 2012. The PduQ enzyme is an alcohol dehydrogenase used to recycle NAD internally within the Pdu microcompartment of Salmonella enterica. PLoS One 7:e47144. doi:10 .1371/journal.pone.0047144.
    • (2012) PLoS One , vol.7
    • Cheng, S.1    Fand, C.2    Sinha, S.3    Bobik, T.A.4


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