메뉴 건너뛰기




Volumn 39, Issue 8, 2010, Pages 1251-1259

Dynamics of strand passage catalyzed by topoisomerase II

Author keywords

Chemomechanical coupling mechanism; DNA topology; DNA transport; Gyrase; Model; Molecular machine

Indexed keywords

ADENOSINE DIPHOSPHATE; ADENOSINE TRIPHOSPHATE; DNA; DNA TOPOISOMERASE (ATP HYDROLYSING);

EID: 77955663856     PISSN: 01757571     EISSN: None     Source Type: Journal    
DOI: 10.1007/s00249-010-0578-y     Document Type: Article
Times cited : (6)

References (39)
  • 1
    • 0027419771 scopus 로고
    • The 43-kilodalton N-terminal fragment of the DNA gyrase B protein hydrolyzes ATP and binds coumarin drugs
    • 10.1021/bi00061a033 1:CAS:528:DyaK3sXhtF2lt74%3D 8383523
    • JA Ali AP Jackson AJ Howells A Maxwell 1993 The 43-kilodalton N-terminal fragment of the DNA gyrase B protein hydrolyzes ATP and binds coumarin drugs Biochemistry 32 2717 2724 10.1021/bi00061a033 1:CAS:528:DyaK3sXhtF2lt74%3D 8383523
    • (1993) Biochemistry , vol.32 , pp. 2717-2724
    • Ali, J.A.1    Jackson, A.P.2    Howells, A.J.3    Maxwell, A.4
  • 2
    • 0029166885 scopus 로고
    • Nucleotide binding to the 43-kilodalton N-terminal fragment of the DNA gyrase B protein
    • 10.1021/bi00030a018 1:CAS:528:DyaK2MXmvVegs70%3D 7626649
    • JA Ali G Orphanides A Maxwell 1995 Nucleotide binding to the 43-kilodalton N-terminal fragment of the DNA gyrase B protein Biochemistry 34 9801 9808 10.1021/bi00030a018 1:CAS:528:DyaK2MXmvVegs70%3D 7626649
    • (1995) Biochemistry , vol.34 , pp. 9801-9808
    • Ali, J.A.1    Orphanides, G.2    Maxwell, A.3
  • 3
    • 0033598716 scopus 로고    scopus 로고
    • Topoisomerase II drives DNA transport by hydrolyzing one ATP
    • 10.1073/pnas.96.24.13685 1:CAS:528:DyaK1MXns1Oqsbo%3D 10570133
    • CL Baird TT Harkins SK Morris JE Lindsley 1999 Topoisomerase II drives DNA transport by hydrolyzing one ATP Proc Natl Acad Sci USA 96 13685 13690 10.1073/pnas.96.24.13685 1:CAS:528:DyaK1MXns1Oqsbo%3D 10570133
    • (1999) Proc Natl Acad Sci USA , vol.96 , pp. 13685-13690
    • Baird, C.L.1    Harkins, T.T.2    Morris, S.K.3    Lindsley, J.E.4
  • 4
    • 34447323073 scopus 로고    scopus 로고
    • Energy coupling in type II topoisomerases: Why do they hydrolyze ATP?
    • DOI 10.1021/bi700789g
    • AD Bates A Maxwell 2007 Energy coupling in type II topoisomerases: why do they hydrolyze ATP? Biochemistry 46 7929 7941 10.1021/bi700789g 1:CAS:528:DC%2BD2sXms12gtL4%3D 17580973 (Pubitemid 47053858)
    • (2007) Biochemistry , vol.46 , Issue.27 , pp. 7929-7941
    • Bates, A.D.1    Maxwell, A.2
  • 5
    • 0030063743 scopus 로고    scopus 로고
    • Energy coupling in Escherichia coli DNA gyrase: The relationship between nucleotide binding, strand passage, and DNA supercoiling
    • DOI 10.1021/bi952433y
    • AD Bates MH O'Dea M Gellert 1996 Energy coupling in Escherichia coli DNA gyrase: the relationship between nucleotide binding, strand passage, and DNA supercoiling Biochemistry 35 1408 1416 10.1021/bi952433y 1:CAS:528: DyaK28XjvVOhsQ%3D%3D 8634270 (Pubitemid 26055276)
    • (1996) Biochemistry , vol.35 , Issue.5 , pp. 1408-1416
    • Bates, A.D.1    O'Dea, M.H.2    Gellert, M.3
  • 6
    • 0029911698 scopus 로고    scopus 로고
    • Recent developments in DNA topoisomerase II structure and mechanism
    • DOI 10.1016/S0959-440X(96)80099-6
    • JM Berger JC Wang 1996 Recent developments in DNA topoisomerase II structure and mechanism Curr Opin Struct Biol 6 84 90 10.1016/S0959-440X(96) 80099-6 1:CAS:528:DyaK28Xhtlamt7Y%3D 8696977 (Pubitemid 26073948)
    • (1996) Current Opinion in Structural Biology , vol.6 , Issue.1 , pp. 84-90
    • Berger, J.M.1    Wang, J.C.2
  • 7
    • 0030045003 scopus 로고    scopus 로고
    • Structure and mechanism of DNA topoisomerase II
    • DOI 10.1038/379225a0
    • JM Berger SJ Gamblin SC Harrison JC Wang 1996 Structure and mechanism of DNA topoisomerase II Nature 379 225 232 10.1038/379225a0 1:CAS:528: DyaK28Xmtl2gsg%3D%3D 8538787 (Pubitemid 26025206)
    • (1996) Nature , vol.379 , Issue.6562 , pp. 225-232
    • Berger, J.M.1    Gamblin, S.J.2    Harrison, S.C.3    Wang, J.C.4
  • 8
    • 0034737716 scopus 로고    scopus 로고
    • Dimerization of Escherichia coli DNA-gyrase B provides a structural mechanism for activating the ATPase catalytic center
    • DOI 10.1074/jbc.275.13.9468
    • L Brino A Urzhumtsev M Mousli C Bronner A Mitschler P Oudet D Moras 2000 Dimerization of Escherichia coli DNA-gyrase B provides a structural mechanism for activating the ATPase catalytic center J Biol Chem 275 9468 9475 10.1074/jbc.275.13.9468 1:CAS:528:DC%2BD3cXitlCru78%3D 10734094 (Pubitemid 30185175)
    • (2000) Journal of Biological Chemistry , vol.275 , Issue.13 , pp. 9468-9475
    • Brino, L.1    Urzhumtsev, A.2    Mousli, M.3    Bronner, C.4    Mitschler, A.5    Oudet, P.6    Moras, D.7
  • 9
    • 0032190561 scopus 로고    scopus 로고
    • Mechanism of action of eukaryotic topoisomerase II and drugs targeted to the enzyme
    • 1:CAS:528:DyaK1cXmtFWisrg%3D 9748545
    • DA Burden N Osheroff 1998 Mechanism of action of eukaryotic topoisomerase II and drugs targeted to the enzyme Biochim Biophys Acta 1400 139 154 1:CAS:528:DyaK1cXmtFWisrg%3D 9748545
    • (1998) Biochim Biophys Acta , vol.1400 , pp. 139-154
    • Burden, D.A.1    Osheroff, N.2
  • 10
    • 0034923502 scopus 로고    scopus 로고
    • DNA topoisomerases: Structure, function, and mechanism
    • DOI 10.1146/annurev.biochem.70.1.369
    • JJ Champoux 2001 DNA topoisomerases: structure, function, and mechanism Annu Rev Biochem 70 369 413 10.1146/annurev.biochem.70.1.369 1:CAS:528:DC%2BD3MXlsVeht7Y%3D 11395412 (Pubitemid 32662215)
    • (2001) Annual Review of Biochemistry , vol.70 , pp. 369-413
    • Champoux, J.J.1
  • 11
    • 0141591551 scopus 로고    scopus 로고
    • Structure of the topoisomerase II ATPase region and its mechanism of inhibition by the chemotherapeutic agent ICRF-187
    • DOI 10.1073/pnas.1832879100
    • S Classen S Olland JM Berger 2003 Structure of the topoisomerase II ATPase region and its mechanism of inhibition by the chemotherapeutic agent ICRF-187 Proc Natl Acad Sci USA 100 10629 10634 10.1073/pnas.1832879100 1:CAS:528:DC%2BD3sXnslynsL8%3D 12963818 (Pubitemid 37140080)
    • (2003) Proceedings of the National Academy of Sciences of the United States of America , vol.100 , Issue.19 , pp. 10629-10634
    • Classen, S.1    Olland, S.2    Berger, J.M.3
  • 12
    • 63349107871 scopus 로고    scopus 로고
    • Analysis of the eukaryotic topoisomerase II DNA gate: A single-molecule FRET and structural perspective
    • 10.1093/nar/gkn1059 1:CAS:528:DC%2BD1MXisFekt70%3D 19155278
    • TRL Collins GG Hammes T-S Hsieh 2009 Analysis of the eukaryotic topoisomerase II DNA gate: a single-molecule FRET and structural perspective Nucleic Acids Res 37 712 720 10.1093/nar/gkn1059 1:CAS:528:DC%2BD1MXisFekt70%3D 19155278
    • (2009) Nucleic Acids Res , vol.37 , pp. 712-720
    • Collins, T.R.L.1    Hammes, G.G.2    Hsieh, T.-S.3
  • 13
    • 2442599792 scopus 로고    scopus 로고
    • Structure, molecular mechanisms, and evolutionary relationships in DNA topoisomerases
    • DOI 10.1146/annurev.biophys.33.110502.140357
    • KD Corbett JM Berger 2004 Structure, molecular mechanisms, and evolutionary relationships in DNA topoisomerases Annu Rev Biophys Biomol Struct 33 95 118 10.1146/annurev.biophys.33.110502.140357 1:CAS:528: DC%2BD2cXltlKktr0%3D 15139806 (Pubitemid 38829953)
    • (2004) Annual Review of Biophysics and Biomolecular Structure , vol.33 , pp. 95-118
    • Corbett, K.D.1    Berger, J.M.2
  • 14
    • 37549023863 scopus 로고    scopus 로고
    • Structural basis for gate-DNA recognition and bending by type IIA topoisomerases
    • 10.1038/nature06396 1:CAS:528:DC%2BD2sXhsVGjsL%2FN 18097402
    • KC Dong JM Berger 2007 Structural basis for gate-DNA recognition and bending by type IIA topoisomerases Nature 450 1201 1205 10.1038/nature06396 1:CAS:528:DC%2BD2sXhsVGjsL%2FN 18097402
    • (2007) Nature , vol.450 , pp. 1201-1205
    • Dong, K.C.1    Berger, J.M.2
  • 15
    • 0032947158 scopus 로고    scopus 로고
    • Quaternary changes in topoisomerase II may direct orthogonal movement of two DNA strands
    • DOI 10.1038/7556
    • D Fass CE Bogden JM Berger 1999 Quaternary changes in topoisomerase II may direct orthogonal movement of two DNA strands Nat Struct Biol 6 322 326 10.1038/7556 1:CAS:528:DyaK1MXisVChsLc%3D 10201398 (Pubitemid 29166478)
    • (1999) Nature Structural Biology , vol.6 , Issue.4 , pp. 322-326
    • Pass, D.1    Bogden, C.E.2    Berger, J.M.3
  • 17
    • 0032387875 scopus 로고    scopus 로고
    • The N-terminal domain of human topoisomerase IIα is a DNA-dependent ATPase
    • DOI 10.1021/bi9818321
    • LP Gardiner DI Roper TR Hammonds A Maxwell 1998 The N-terminal domain of human topoisomerase II-alpha is a DNA-dependent ATPase Biochemistry 37 16997 17004 10.1021/bi9818321 1:CAS:528:DyaK1cXntFegt7g%3D 9836594 (Pubitemid 28566778)
    • (1998) Biochemistry , vol.37 , Issue.48 , pp. 16997-17004
    • Gardiner, L.P.1    Roper, D.I.2    Hammonds, T.R.3    Maxwell, A.4
  • 18
    • 0032546589 scopus 로고    scopus 로고
    • Pre-steady-state analysis of ATP hydrolysis by Saccharomyces cerevisiae DNA topoisomerase II. 1. A DNA-dependent burst in ATP hydrolysis
    • DOI 10.1021/bi9729099
    • TT Harkins JE Lindsley 1998 Pre-steady-state analysis of ATP hydrolysis by saccharomyces cerevisiae DNA topoisomerase II. 1. A DNA-dependent burst in ATP hydrolysis Biochemistry 37 7292 7298 10.1021/bi9729099 1:CAS:528: DyaK1cXislGgur4%3D 9585543 (Pubitemid 28235212)
    • (1998) Biochemistry , vol.37 , Issue.20 , pp. 7292-7298
    • Harkins, T.T.1    Lindsley, J.E.2
  • 19
    • 0032546611 scopus 로고    scopus 로고
    • Pre-steady-state analysis of ATP hydrolysis by Saccharomyces cerevisiae DNA topoisomerase II. 2. Kinetic mechanism for the sequential hydrolysis of two ATP
    • DOI 10.1021/bi9729108
    • TT Harkins TJ Lewis JE Lindsley 1998 Pre-steady-state analysis of ATP hydrolysis by saccharomyces cerevisiae DNA topoisomerase II. 2. Kinetic mechanism for the sequential hydrolysis of two ATP Biochemistry 37 7299 7312 10.1021/bi9729108 1:CAS:528:DyaK1cXislGgur8%3D 9585544 (Pubitemid 28235213)
    • (1998) Biochemistry , vol.37 , Issue.20 , pp. 7299-7312
    • Harkins, T.T.1    Lewis, T.J.2    Lindsley, J.E.3
  • 20
    • 0037155222 scopus 로고    scopus 로고
    • ATPase domain of eukaryotic DNA topoisomerase II. Inhibition of ATPase activity by the anti-cancer drug bisdioxopiperazine and ATP/ADP-induced dimerization
    • DOI 10.1074/jbc.M111394200
    • T Hu H Sage TS Hsieh 2002 ATPase domain of eukaryotic DNA topoisomerase II J Biol Chem 277 5944 5951 10.1074/jbc.M111394200 1:CAS:528: DC%2BD38XhvFyrtrg%3D 11850431 (Pubitemid 34968375)
    • (2002) Journal of Biological Chemistry , vol.277 , Issue.8 , pp. 5944-5951
    • Hu, T.1    Sage, H.2    Hsieh, T.-S.3
  • 21
    • 0037166285 scopus 로고    scopus 로고
    • An open conformation of the Thermus thermophilus gyrase B ATP-binding domain
    • DOI 10.1074/jbc.M111740200
    • V Lamour L Hoermann JM Jeltsch P Oudet D Moras 2002 An open conformation of the Thermus thermophilus gyrase B ATP-binding domain J Biol Chem 277 18947 18953 10.1074/jbc.M111740200 1:CAS:528:DC%2BD38XktFCht74%3D 11850422 (Pubitemid 34952456)
    • (2002) Journal of Biological Chemistry , vol.277 , Issue.21 , pp. 18947-18953
    • Lamour, V.1    Hoermann, L.2    Jeltsch, J.-M.3    Oudet, P.4    Moras, D.5
  • 22
    • 0027462987 scopus 로고
    • Study of allosteric communication between protomers by immunotagging
    • DOI 10.1038/361749a0
    • JE Lindsley JC Wang 1993 Study of allosteric communication between protomers by immunotagging Nature 361 749 750 10.1038/361749a0 1:CAS:528:DyaK3sXhvVKrtb8%3D 7680110 (Pubitemid 23070487)
    • (1993) Nature , vol.361 , Issue.6414 , pp. 749-750
    • Lindsley, J.E.1    Wang, J.C.2
  • 23
    • 0000301456 scopus 로고    scopus 로고
    • Crystal structure of the breakage-reunion domain of DNA gyrase
    • DOI 10.1038/42294
    • JH Morais Cabral AP Jackson CV Smith N Shikotra A Maxwell RC Liddington 1997 Crystal structure of the break age-reunion domain of DNA gyrase Nature 388 903 906 10.1038/42294 1:CAS:528:DyaK2sXlslansL0%3D 9278055 (Pubitemid 27377066)
    • (1997) Nature , vol.388 , Issue.6645 , pp. 903-906
    • Morais Cabral, J.H.1    Jackson, A.P.2    Smith, C.V.3    Shikotra, N.4    Maxwell, A.5    Liddington, R.C.6
  • 24
    • 47749137356 scopus 로고    scopus 로고
    • Coupling between ATP binding and DNA cleavage by DNA topoisomerase II: A unified kinetic and structural mechanism
    • 10.1074/jbc.M710014200
    • F Mueller D Herschalg 2008 Coupling between ATP binding and DNA cleavage by DNA topoisomerase II: a unified kinetic and structural mechanism J Biol Chem 283 17463 17476 10.1074/jbc.M710014200
    • (2008) J Biol Chem , vol.283 , pp. 17463-17476
    • Mueller, F.1    Herschalg, D.2
  • 25
    • 0033618406 scopus 로고    scopus 로고
    • 2-terminal fragments of yeast DNA topoisomerase II
    • DOI 10.1074/jbc.274.31.21688
    • 2-terminal fragments of yeast DNA topoisomerase II J Biol Chem 274 21688 21694 10.1074/jbc.274.31.21688 1:CAS:528:DyaK1MXltVymsLg%3D 10419479 (Pubitemid 29359994)
    • (1999) Journal of Biological Chemistry , vol.274 , Issue.31 , pp. 21688-21694
    • Olland, S.1    Wang, J.C.2
  • 26
    • 63349108302 scopus 로고    scopus 로고
    • Topoisomerase II: A fitted mechanism for the chromatin landscape
    • 10.1093/nar/gkn994 1:CAS:528:DC%2BD1MXisFekurY%3D 19059997
    • J Roca 2009 Topoisomerase II: a fitted mechanism for the chromatin landscape Nucleic Acids Res 37 721 730 10.1093/nar/gkn994 1:CAS:528: DC%2BD1MXisFekurY%3D 19059997
    • (2009) Nucleic Acids Res , vol.37 , pp. 721-730
    • Roca, J.1
  • 27
    • 0026448670 scopus 로고
    • The capture of a DNA double helix by an ATP-dependent protein clamp: A key step in DNA transport by type II DNA topoisomerases
    • 10.1016/0092-8674(92)90558-T 1:CAS:528:DyaK3sXhtF2lu7k%3D 1330327
    • J Roca JC Wang 1992 The capture of a DNA double helix by an ATP-dependent protein clamp: a key step in DNA transport by type II DNA topoisomerases Cell 71 833 840 10.1016/0092-8674(92)90558-T 1:CAS:528:DyaK3sXhtF2lu7k%3D 1330327
    • (1992) Cell , vol.71 , pp. 833-840
    • Roca, J.1    Wang, J.C.2
  • 28
    • 0028334718 scopus 로고
    • DNA transport by a type II DNA topoisomerase: Evidence in favor of a two- gate mechanism
    • DOI 10.1016/0092-8674(94)90222-4
    • J Roca JC Wang 1994 DNA transport by a type II DNA topoisomerase: evidence in favor of a two-gate mechanism Cell 77 609 616 10.1016/0092-8674(94) 90222-4 1:CAS:528:DyaK2cXltVart7s%3D 8187179 (Pubitemid 24154000)
    • (1994) Cell , vol.77 , Issue.4 , pp. 609-616
    • Roca, J.1    Wang, J.C.2
  • 30
    • 28844455588 scopus 로고    scopus 로고
    • Recent advances in understanding structure-function relationships in the type II topoisomerase mechanism
    • DOI 10.1042/BST20051465
    • AJ Schoeffler JM Berger 2005 Recent advances in understanding structure-function relationships in the type II topoisomerase mechanism Biochem Soc Trans 33 1465 1470 10.1042/BST20051465 1:CAS:528:DC%2BD2MXhtFCqsLjE 16246147 (Pubitemid 41779716)
    • (2005) Biochemical Society Transactions , vol.33 , Issue.6 , pp. 1465-1470
    • Schoeffler, A.J.1    Berger, J.M.2
  • 31
    • 0037458744 scopus 로고    scopus 로고
    • A human topoisomerase IIα heterodimer with only one ATP binding site can go through successive catalytic cycles
    • DOI 10.1074/jbc.M210332200
    • C Skouboe L Bjergbaek VH Oestergaard MK Larsen BR Knudsen AH Andersen 2003 A human topoisomerase II α heterodimer with only one ATP binding site can go through successive catalytic cycles J Biol Chem 278 5768 5774 10.1074/jbc.M210332200 1:CAS:528:DC%2BD3sXht1aqsbg%3D 12480934 (Pubitemid 36800822)
    • (2003) Journal of Biological Chemistry , vol.278 , Issue.8 , pp. 5768-5774
    • Skouboe, C.1    Bjergbaek, L.2    Oestergaard, V.H.3    Larsen, M.K.4    Knudsen, B.R.5    Andersen, A.H.6
  • 34
    • 0030014783 scopus 로고    scopus 로고
    • DNA topoisomerases
    • 10.1146/annurev.bi.65.070196.003223 1:CAS:528:DyaK28XktFamur0%3D 8811192
    • JC Wang 1996 DNA topoisomerases Annu Rev Biochem 65 635 692 10.1146/annurev.bi.65.070196.003223 1:CAS:528:DyaK28XktFamur0%3D 8811192
    • (1996) Annu Rev Biochem , vol.65 , pp. 635-692
    • Wang, J.C.1
  • 35
    • 27744591551 scopus 로고    scopus 로고
    • Nucleotide-dependent domain movement in the ATPase domain of a human type IIA DNA topoisomerase
    • DOI 10.1074/jbc.M506520200
    • H Wei AJ Ruthenburg SK Bechis GL Verdine 2005 Nucleotide-dependent domain movement in the ATPase domain of a human type IIA DNA topoisomerase J Biol Chem 280 37041 37047 10.1074/jbc.M506520200 1:CAS:528:DC%2BD2MXhtFKhtLrO 16100112 (Pubitemid 41587788)
    • (2005) Journal of Biological Chemistry , vol.280 , Issue.44 , pp. 37041-37047
    • Wei, H.1    Ruthenburg, A.J.2    Bechis, S.K.3    Verdine, G.L.4
  • 36
    • 0031879578 scopus 로고    scopus 로고
    • Trapping and wiggling: Elastohydrodynamics of driven microfilaments
    • 10.1016/S0006-3495(98)74029-9 1:CAS:528:DyaK1cXhs1KrsrY%3D 9533717
    • CH Wiggins D Riveline A Ott RE Goldstein 1998 Trapping and wiggling: elastohydrodynamics of driven microfilaments Biophys J 74 1043 1060 10.1016/S0006-3495(98)74029-9 1:CAS:528:DyaK1cXhs1KrsrY%3D 9533717
    • (1998) Biophys J , vol.74 , pp. 1043-1060
    • Wiggins, C.H.1    Riveline, D.2    Ott, A.3    Goldstein, R.E.4
  • 37
    • 0026428621 scopus 로고
    • Crystal structure of an N-terminal fragment of the DNA gyrase B protein
    • 10.1038/351624a0 1:CAS:528:DyaK3MXltV2gs7o%3D 1646964
    • DB Wigley GJ Davies EJ Dodson A Maxwell G Dodson 1991 Crystal structure of an N-terminal fragment of the DNA gyrase B protein Nature 351 624 629 10.1038/351624a0 1:CAS:528:DyaK3MXltV2gs7o%3D 1646964
    • (1991) Nature , vol.351 , pp. 624-629
    • Wigley, D.B.1    Davies, G.J.2    Dodson, E.J.3    Maxwell, A.4    Dodson, G.5
  • 38
    • 49349091072 scopus 로고    scopus 로고
    • Stepping behavior of two-headed kinesin motors
    • 10.1016/j.bbabio.2008.04.040 1:CAS:528:DC%2BD1cXpvF2quro%3D 18501186
    • P Xie 2008 Stepping behavior of two-headed kinesin motors Biochim Biophys Acta 1777 1195 1202 10.1016/j.bbabio.2008.04.040 1:CAS:528:DC%2BD1cXpvF2quro%3D 18501186
    • (2008) Biochim Biophys Acta , vol.1777 , pp. 1195-1202
    • Xie, P.1
  • 39
    • 36549042972 scopus 로고    scopus 로고
    • Processivity of single-headed kinesin motors
    • DOI 10.1016/j.bbabio.2007.09.006, PII S0005272807002241
    • P Xie S-X Dou P-Y Wang 2007 Processivity of single-headed kinesin motors Biochim Biophys Acta 1767 1418 1427 10.1016/j.bbabio.2007.09.006 1:CAS:528:DC%2BD2sXhtlyhsrbF 17976515 (Pubitemid 350179712)
    • (2007) Biochimica et Biophysica Acta - Bioenergetics , vol.1767 , Issue.12 , pp. 1418-1427
    • Xie, P.1    Dou, S.-X.2    Wang, P.-Y.3


* 이 정보는 Elsevier사의 SCOPUS DB에서 KISTI가 분석하여 추출한 것입니다.