메뉴 건너뛰기




Volumn 11, Issue 10, 2004, Pages 936-944

Intron-encoded homing endonuclease I-TevI also functions as a transcriptional autorepressor

Author keywords

[No Author keywords available]

Indexed keywords

ENDONUCLEASE; ENDONUCLEASE I TEVI; UNCLASSIFIED DRUG;

EID: 4744359807     PISSN: 15459993     EISSN: None     Source Type: Journal    
DOI: 10.1038/nsmb823     Document Type: Article
Times cited : (36)

References (49)
  • 1
    • 0029059329 scopus 로고
    • Prokaryotic introns and inteins: A panoply of form and function
    • Belfort, M., Reaban, M.E., Coetzee, T. & Dalgaard, J.Z. Prokaryotic introns and inteins: a panoply of form and function. J. Bacteriol. 177, 3897-3903 (1995).
    • (1995) J. Bacteriol. , vol.177 , pp. 3897-3903
    • Belfort, M.1    Reaban, M.E.2    Coetzee, T.3    Dalgaard, J.Z.4
  • 3
    • 85158033913 scopus 로고    scopus 로고
    • Mobile introns: Pathways and proteins
    • (eds. Craig, N., Craigie, R., Gellert, M. & Lambowitz, A.) (ASM Press, Washington DC)
    • Belfort, M., Derbyshire, V., Cousineau, B. & Lambowitz, A. Mobile introns: pathways and proteins. In Mobile DNA II (eds. Craig, N., Craigie, R., Gellert, M. & Lambowitz, A.) 761-783 (ASM Press, Washington DC, 2002).
    • (2002) Mobile DNA II , pp. 761-783
    • Belfort, M.1    Derbyshire, V.2    Cousineau, B.3    Lambowitz, A.4
  • 4
    • 0024418461 scopus 로고
    • Mobile introns: Definition of terms and recommended nomenclature
    • Dujon, B. et al. Mobile introns: definition of terms and recommended nomenclature. Gene 82, 115-118 (1989).
    • (1989) Gene , vol.82 , pp. 115-118
    • Dujon, B.1
  • 5
    • 0030044828 scopus 로고    scopus 로고
    • Intron mobility in phage T4 occurs in the context of recombination- dependent DNA replication by way of multiple pathways
    • Mueller, J.E., Clyman, J., Huang, Y.-J., Parker, M.M. & Belfort, M. Intron mobility in phage T4 occurs in the context of recombination-dependent DNA replication by way of multiple pathways. Genes Dev. 10, 351-364 (1996).
    • (1996) Genes Dev. , vol.10 , pp. 351-364
    • Mueller, J.E.1    Clyman, J.2    Huang, Y.-J.3    Parker, M.M.4    Belfort, M.5
  • 6
    • 0024346261 scopus 로고
    • Bacteriophage introns: Parasites within parasites?
    • Belfort, M. Bacteriophage introns: parasites within parasites? Trends Genet. 5, 209-213 (1989).
    • (1989) Trends Genet. , vol.5 , pp. 209-213
    • Belfort, M.1
  • 7
    • 0037180394 scopus 로고    scopus 로고
    • Catalytic mechanisms of restriction and homing endonucleases
    • Galburt, E.A. & Stoddard, B.L. Catalytic mechanisms of restriction and homing endonucleases. Biochemistry 41, 13851-13860 (2002).
    • (2002) Biochemistry , vol.41 , pp. 13851-13860
    • Galburt, E.A.1    Stoddard, B.L.2
  • 8
    • 0033567272 scopus 로고    scopus 로고
    • Homing endonucleases: Structure, function and evolution
    • Jurica, M.S. & Stoddard, B.L. Homing endonucleases: structure, function and evolution. Cell. Mol. Life Sci. 55, 1304-1326 (1999).
    • (1999) Cell. Mol. Life Sci. , vol.55 , pp. 1304-1326
    • Jurica, M.S.1    Stoddard, B.L.2
  • 9
    • 0027207339 scopus 로고
    • The td intron endonuclease makes extensive sequence tolerant contacts across the minor groove of its DNA target
    • Bryk, M. et al. The td intron endonuclease makes extensive sequence tolerant contacts across the minor groove of its DNA target. EMBO J. 12, 2141-2149 (1993).
    • (1993) EMBO J. , vol.12 , pp. 2141-2149
    • Bryk, M.1
  • 10
    • 0028943142 scopus 로고
    • Selection of a remote cleavage site by I-TevI, the td intron-encoded endonuclease
    • Bryk, M., Belisle, M., Mueller, J.E. & Belfort, M. Selection of a remote cleavage site by I-TevI, the td intron-encoded endonuclease. J. Mol. Biol. 247, 197-210 (1995).
    • (1995) J. Mol. Biol. , vol.247 , pp. 197-210
    • Bryk, M.1    Belisle, M.2    Mueller, J.E.3    Belfort, M.4
  • 11
    • 0035883706 scopus 로고    scopus 로고
    • Homing endonucleases: Structural and functional insight into the catalysis of intron/intein mobility
    • Chevalier, B.S. & Stoddard, B.L. Homing endonucleases: structural and functional insight into the catalysis of intron/intein mobility. Nucleic Acids Res. 29, 3757-3774 (2001).
    • (2001) Nucleic Acids Res. , vol.29 , pp. 3757-3774
    • Chevalier, B.S.1    Stoddard, B.L.2
  • 12
    • 0028113405 scopus 로고
    • An intron in the thymidylate synthase gene of Bacillus bacteriophage β22: Evidence for independent evolution of a gene, its group I intron, and the intron open reading frame
    • Bechhofer, D.H., Hue, K.K. & Shub, D.A. An intron in the thymidylate synthase gene of Bacillus bacteriophage β22: Evidence for independent evolution of a gene, its group I intron, and the intron open reading frame. Proc. Natl. Acad. Sci. USA 91, 11669-11673 (1994).
    • (1994) Proc. Natl. Acad. Sci. USA , vol.91 , pp. 11669-11673
    • Bechhofer, D.H.1    Hue, K.K.2    Shub, D.A.3
  • 13
    • 0034940222 scopus 로고    scopus 로고
    • Related homing endonucleases I-Bmol and I-TevI use different strategies to cleave homologous recognition sites
    • Edgell, D.R. & Shub, D.A. Related homing endonucleases I-Bmol and I-TevI use different strategies to cleave homologous recognition sites. Proc. Natl. Acad. Sci. USA 98, 7898-7903 (2001).
    • (2001) Proc. Natl. Acad. Sci. USA , vol.98 , pp. 7898-7903
    • Edgell, D.R.1    Shub, D.A.2
  • 14
    • 0001423441 scopus 로고
    • Intervening sequence in the thymidylate synthase gene of bacteriophage T4
    • Chu, F.K., Maley, G.F., Maley, F. & Belfort, M. Intervening sequence in the thymidylate synthase gene of bacteriophage T4. Proc. Natl. Acad. Sci. USA 81, 3049-3053 (1984).
    • (1984) Proc. Natl. Acad. Sci. USA , vol.81 , pp. 3049-3053
    • Chu, F.K.1    Maley, G.F.2    Maley, F.3    Belfort, M.4
  • 15
    • 0023046266 scopus 로고
    • Multiple self-splicing introns in bacteriophage T4: Evidence from autocatalytic GTP labeling of RNA in vitro
    • Gott, J.M., Shub, D.A. & Belfort, M. Multiple self-splicing introns in bacteriophage T4: Evidence from autocatalytic GTP labeling of RNA in vitro. Cell 47, 81-87 (1986).
    • (1986) Cell , vol.47 , pp. 81-87
    • Gott, J.M.1    Shub, D.A.2    Belfort, M.3
  • 16
    • 0028050514 scopus 로고
    • Intron-containing T4 bacteriophage gene sunY encodes an anaerobic ribonucleotide reductase
    • Young, P., Ohman, M., Xu, M.Q., Shub, D.A. & Sjoberg, B.-M. Intron-containing T4 bacteriophage gene sunY encodes an anaerobic ribonucleotide reductase. J. Biol. Chem. 269, 20229-20232 (1994).
    • (1994) J. Biol. Chem. , vol.269 , pp. 20229-20232
    • Young, P.1    Ohman, M.2    Xu, M.Q.3    Shub, D.A.4    Sjoberg, B.-M.5
  • 17
    • 0024965995 scopus 로고
    • Intron mobility in the T-even phages: High frequency inheritance of group I introns promoted by intron open reading frames
    • Quirk, S.M., Bell-Pedersen, D. & Belfort, M. Intron mobility in the T-even phages: high frequency inheritance of group I introns promoted by intron open reading frames. Cell 56, 455-465 (1989).
    • (1989) Cell , vol.56 , pp. 455-465
    • Quirk, S.M.1    Bell-Pedersen, D.2    Belfort, M.3
  • 18
    • 0025796852 scopus 로고
    • The phage T4 nrdB intron: A deletion mutant of a version found in the wild
    • Eddy, S.R. & Gold, L. The phage T4 nrdB intron: a deletion mutant of a version found in the wild. Genes Dev. 5, 1032-1041 (1991).
    • (1991) Genes Dev. , vol.5 , pp. 1032-1041
    • Eddy, S.R.1    Gold, L.2
  • 19
    • 0024243749 scopus 로고
    • Genes within genes: Independent expression of phage T4 intron open reading frames and the genes in which they reside
    • Gott, J.M. et al. Genes within genes: Independent expression of phage T4 intron open reading frames and the genes in which they reside. Genes Dev. 2, 1791-1799 (1988).
    • (1988) Genes Dev. , vol.2 , pp. 1791-1799
    • Gott, J.M.1
  • 20
    • 0032079494 scopus 로고    scopus 로고
    • A ribosomal function is necessary for efficient splicing of the T4 phage thymidylatesynthase intron in vivo
    • Semrad, K. & Schroeder, R. A ribosomal function is necessary for efficient splicing of the T4 phage thymidylatesynthase intron in vivo. Genes Dev. 12, 1327-1337 (1998).
    • (1998) Genes Dev. , vol.12 , pp. 1327-1337
    • Semrad, K.1    Schroeder, R.2
  • 21
    • 0035898540 scopus 로고    scopus 로고
    • Intertwined structure of the DNA-binding domain of intron endonuclease I-TevI with its substrate
    • Van Roey, P., Waddling, C.A., Rox, K.M., Belfort, M. & Derbyshire, V. Intertwined structure of the DNA-binding domain of intron endonuclease I-TevI with its substrate. EMBO J. 20, 3631-3637 (2001).
    • (2001) EMBO J. , vol.20 , pp. 3631-3637
    • Van Roey, P.1    Waddling, C.A.2    Rox, K.M.3    Belfort, M.4    Derbyshire, V.5
  • 22
    • 0029962990 scopus 로고    scopus 로고
    • The intron-encoded endonuclease I-TevI I binds across the minor groove and induces two distinct conformational changes in its DNA substrate
    • Loizos, N., Silva, G.H. & Belfort, M. The intron-encoded endonuclease I-TevI I binds across the minor groove and induces two distinct conformational changes in its DNA substrate. J. Mol. Biol. 255, 412-424 (1996).
    • (1996) J. Mol. Biol. , vol.255 , pp. 412-424
    • Loizos, N.1    Silva, G.H.2    Belfort, M.3
  • 23
    • 0031556945 scopus 로고    scopus 로고
    • Two-domain structure of the td intron-encoded endonuclease I-TevI correlates with the two-domain configuration of the homing site
    • Derbyshire, V., Kowalski, J.C., Dansereau, J.T., Hauer, C.R. & Belfort, M. Two-domain structure of the td intron-encoded endonuclease I-TevI correlates with the two-domain configuration of the homing site. J. Mol. Biol. 265, 494-506 (1997).
    • (1997) J. Mol. Biol. , vol.265 , pp. 494-506
    • Derbyshire, V.1    Kowalski, J.C.2    Dansereau, J.T.3    Hauer, C.R.4    Belfort, M.5
  • 24
    • 0037172989 scopus 로고    scopus 로고
    • Zinc finger as distance determinant in the flexible linker of intron endonuclease I-TevI
    • Dean, A.B. et al. Zinc finger as distance determinant in the flexible linker of intron endonuclease I-TevI. Proc. Natl. Acad. Sci. USA 99, 8554-8561 (2002).
    • (2002) Proc. Natl. Acad. Sci. USA , vol.99 , pp. 8554-8561
    • Dean, A.B.1
  • 25
    • 0042706146 scopus 로고    scopus 로고
    • Moonlighting proteins: Old proteins learning new tricks
    • Jeffery, C.J. Moonlighting proteins: old proteins learning new tricks. Trends Genet. 19, 415-417 (2003).
    • (2003) Trends Genet. , vol.19 , pp. 415-417
    • Jeffery, C.J.1
  • 26
    • 0346123001 scopus 로고    scopus 로고
    • Two for the price of one: A bifunctional intron-encoded DNA endonuclease-RNA maturase
    • Belfort, M. Two for the price of one: a bifunctional intron-encoded DNA endonuclease-RNA maturase. Genes Dev. 17, 2860-2863 (2003).
    • (2003) Genes Dev. , vol.17 , pp. 2860-2863
    • Belfort, M.1
  • 27
    • 0346030573 scopus 로고    scopus 로고
    • Structural and biochemical analyses of DNA and RNA binding by a bifunctional homing endonuclease and group I intron splicing factor
    • Bolduc, J.M. et al. Structural and biochemical analyses of DNA and RNA binding by a bifunctional homing endonuclease and group I intron splicing factor. Genes Dev. 17, 2875-2888 (2003).
    • (2003) Genes Dev. , vol.17 , pp. 2875-2888
    • Bolduc, J.M.1
  • 28
    • 0141925715 scopus 로고    scopus 로고
    • Involvement of the VDE homing endonuclease and rapamycin in regulation of the Saccharomyces cerevisiae GSH11 gene encoding the high affinity glutathione transporter
    • Miyake, T, Hiraishi, H., Sammoto, H. & Ono, B.-I. Involvement of the VDE homing endonuclease and rapamycin in regulation of the Saccharomyces cerevisiae GSH11 gene encoding the high affinity glutathione transporter. J. Biol. Chem. 278, 39632-39636 (2003).
    • (2003) J. Biol. Chem. , vol.278 , pp. 39632-39636
    • Miyake, T.1    Hiraishi, H.2    Sammoto, H.3    Ono, B.-I.4
  • 29
    • 0027973379 scopus 로고
    • Evolution of mobile group I introns: Recognition of intron sequences by an intron-encoded endonuclease
    • Loizos, N., Tillier, E.R.M. & Belfort, M. Evolution of mobile group I introns: Recognition of intron sequences by an intron-encoded endonuclease. Proc. Natl. Acad. Sci. USA 91, 11983-11987 (1994).
    • (1994) Proc. Natl. Acad. Sci. USA , vol.91 , pp. 11983-11987
    • Loizos, N.1    Tillier, E.R.M.2    Belfort, M.3
  • 30
    • 0035902549 scopus 로고    scopus 로고
    • The tight linkage between DNA replication and double-strand break repair in bacteriophage T4
    • George, J.W., Stohr, B.A., Tomso, D.J. & Kreuzer, K.N. The tight linkage between DNA replication and double-strand break repair in bacteriophage T4. Proc. Natl. Acad. Sci. USA 98, 8290-8297 (2001).
    • (2001) Proc. Natl. Acad. Sci. USA , vol.98 , pp. 8290-8297
    • George, J.W.1    Stohr, B.A.2    Tomso, D.J.3    Kreuzer, K.N.4
  • 31
    • 0036468146 scopus 로고    scopus 로고
    • Intronless homing: Site-specific endonuclease SegF of bacteriophage T4 mediates localized marker exclusion analogous to homing endonucleases of group I introns
    • Belle, A., Landthaler, M. & Shub, D.A. Intronless homing: site-specific endonuclease SegF of bacteriophage T4 mediates localized marker exclusion analogous to homing endonucleases of group I introns. Genes Dev. 16, 351-362 (2002).
    • (2002) Genes Dev. , vol.16 , pp. 351-362
    • Belle, A.1    Landthaler, M.2    Shub, D.A.3
  • 32
    • 0037117778 scopus 로고    scopus 로고
    • Selfish DNA new abode for homing endonucleases
    • Edgell, D.R. Selfish DNA: new abode for homing endonucleases. Curr. Biol. 12, R276-R278 (2002).
    • (2002) Curr. Biol. , vol.12
    • Edgell, D.R.1
  • 33
  • 34
    • 0036828959 scopus 로고    scopus 로고
    • Catalytic domain structure and hypothesis for function of GIY-YIG intron endonuclease I-TevI
    • Van Roey, P., Meehan, L., Kowalski, J., Belfort, M. & Derbyshire, V. Catalytic domain structure and hypothesis for function of GIY-YIG intron endonuclease I-TevI. Nat. Struct. Biol. 9, 806-811 (2002).
    • (2002) Nat. Struct. Biol. , vol.9 , pp. 806-811
    • Van Roey, P.1    Meehan, L.2    Kowalski, J.3    Belfort, M.4    Derbyshire, V.5
  • 35
    • 0025883512 scopus 로고
    • I-TevI, the endonuclease encoded by the mobile td intron, recognizes binding and cleavage domains on its DNA target
    • Bell-Pedersen, D., Quirk, S.M., Bryk, M. & Belfort, M. I-TevI, the endonuclease encoded by the mobile td intron, recognizes binding and cleavage domains on its DNA target. Proc. Natl. Acad. Sci. USA 88, 7719-7723 (1991).
    • (1991) Proc. Natl. Acad. Sci. USA , vol.88 , pp. 7719-7723
    • Bell-Pedersen, D.1    Quirk, S.M.2    Bryk, M.3    Belfort, M.4
  • 36
    • 0032185110 scopus 로고    scopus 로고
    • DNA recognition and cleavage by the LAGLIDADG homing endonuclease I-Crel
    • Jurica, M.S., Monnat, R.J. Jr. & Stoddard, B.L. DNA recognition and cleavage by the LAGLIDADG homing endonuclease I-Crel. Mol. Cell 2, 469-476 (1998).
    • (1998) Mol. Cell , vol.2 , pp. 469-476
    • Jurica, M.S.1    Monnat Jr., R.J.2    Stoddard, B.L.3
  • 37
    • 0038061526 scopus 로고    scopus 로고
    • Flexible DNA target site recognition by divergent homing endonuclease isoschizomers I-Crel and I-Msol
    • Chevalier, B., Turmel, M., Lemieux, C., Monnat, R.J.J. & Stoddard, B.L. Flexible DNA target site recognition by divergent homing endonuclease isoschizomers I-Crel and I-Msol. J. Mol. Biol. 329, 253-269 (2003).
    • (2003) J. Mol. Biol. , vol.329 , pp. 253-269
    • Chevalier, B.1    Turmel, M.2    Lemieux, C.3    Monnat, R.J.J.4    Stoddard, B.L.5
  • 38
    • 0032474760 scopus 로고    scopus 로고
    • DNA binding and cleavage by the nuclear intron-encoded homing endonuclease I-Ppol
    • Flick, K.E., Jurica, M.S., Monnat, R.J. Jr. & Stoddard, B.L. DNA binding and cleavage by the nuclear intron-encoded homing endonuclease I-Ppol. Nature 394, 96-101 (1998).
    • (1998) Nature , vol.394 , pp. 96-101
    • Flick, K.E.1    Jurica, M.S.2    Monnat Jr., R.J.3    Stoddard, B.L.4
  • 39
    • 0001619320 scopus 로고
    • On the structure of the glucosylated hydroxymethylcytosine nucleotides of coliphages T2, T4, and T6
    • Lehman, I.R. & Pratt, E.A. On the structure of the glucosylated hydroxymethylcytosine nucleotides of coliphages T2, T4, and T6. J. Biol. Chem. 235, 3254-3259 (1960).
    • (1960) J. Biol. Chem. , vol.235 , pp. 3254-3259
    • Lehman, I.R.1    Pratt, E.A.2
  • 40
    • 0029048413 scopus 로고
    • Structure of BamHI endonuclease bound to DNA: Partial folding and unfolding on DNA binding
    • Newman, M., Strzelecka, T., Dorner, L.F., Schildkraut, I. & Aggarwal, A.K. Structure of BamHI endonuclease bound to DNA: partial folding and unfolding on DNA binding. Science 269, 656-663 (1995).
    • (1995) Science , vol.269 , pp. 656-663
    • Newman, M.1    Strzelecka, T.2    Dorner, L.F.3    Schildkraut, I.4    Aggarwal, A.K.5
  • 41
    • 0033634983 scopus 로고    scopus 로고
    • Structure of BamHI bound to nonspecific DNA: A model for DNA sliding
    • Viadiu, H. & Aggarwal, A.K. Structure of BamHI bound to nonspecific DNA: a model for DNA sliding. Mol. Cell 5, 889-895 (2000).
    • (2000) Mol. Cell , vol.5 , pp. 889-895
    • Viadiu, H.1    Aggarwal, A.K.2
  • 42
    • 0023512003 scopus 로고
    • Genetic delineation of functional components of the group I intron in the phage T4 td gene
    • Belfort, M., Chandry, P.S. & Pedersen-Lane, J. Genetic delineation of functional components of the group I intron in the phage T4 td gene. Cold Spring Harb. Symp. Quant. Biol. 52, 181-192 (1987).
    • (1987) Cold Spring Harb. Symp. Quant. Biol. , vol.52 , pp. 181-192
    • Belfort, M.1    Chandry, P.S.2    Pedersen-Lane, J.3
  • 43
    • 0016616136 scopus 로고
    • Characterization of a new class of deletions of the D region of the bacteriophage T4 genome
    • Depew, R.E., Snopek, T.J. & Cozzarelli, N.R. Characterization of a new class of deletions of the D region of the bacteriophage T4 genome. Virology 64, 144-145 (1975).
    • (1975) Virology , vol.64 , pp. 144-145
    • Depew, R.E.1    Snopek, T.J.2    Cozzarelli, N.R.3
  • 44
    • 0022218185 scopus 로고
    • Bacteriophage T4, a new vector for the expression of cloned genes
    • Shub, D.A. & Casna, N.J. Bacteriophage T4, a new vector for the expression of cloned genes. Gene 37, 31-36 (1985).
    • (1985) Gene , vol.37 , pp. 31-36
    • Shub, D.A.1    Casna, N.J.2
  • 45
    • 0025261344 scopus 로고
    • Application of fluorescence energy transfer and polarization to monitor Escherichia coli cAMP receptor protein and lac promoter interaction
    • Heyduk, T. & Lee, J.C. Application of fluorescence energy transfer and polarization to monitor Escherichia coli cAMP receptor protein and lac promoter interaction. Proc. Natl. Acad. Sci. USA 87, 1744-1748 (1990).
    • (1990) Proc. Natl. Acad. Sci. USA , vol.87 , pp. 1744-1748
    • Heyduk, T.1    Lee, J.C.2
  • 46
    • 0037850968 scopus 로고    scopus 로고
    • Importance of a single base pair for discrimination between intron-containing and intronless alleles by endonuclease I-Bmol
    • Edgell, D. R., Stanger, M.J. & Belfort, M. Importance of a single base pair for discrimination between intron-containing and intronless alleles by endonuclease I-Bmol. Curr. Biol. 13, 973-978 (2003).
    • (2003) Curr. Biol. , vol.13 , pp. 973-978
    • Edgell, D.R.1    Stanger, M.J.2    Belfort, M.3
  • 47
    • 3543012707 scopus 로고    scopus 로고
    • Crystallography & NMR system: A new software suite for macromolecular structure determination
    • Brunger, A.T. et al. Crystallography & NMR system: A new software suite for macromolecular structure determination. Acta Crystallogr. D 54, 905-921 (1998).
    • (1998) Acta Crystallogr. D , vol.54 , pp. 905-921
    • Brunger, A.T.1
  • 48
    • 84889120137 scopus 로고
    • Improved methods for building protein models in electron density maps and the location of errors in these models
    • Jones, T.A., Zou, J.Y., Cowan, S.W. & Kjeldgaard, M. Improved methods for building protein models in electron density maps and the location of errors in these models. Acta Crystallogr. A 47, 110-119 (1991).
    • (1991) Acta Crystallogr. A , vol.47 , pp. 110-119
    • Jones, T.A.1    Zou, J.Y.2    Cowan, S.W.3    Kjeldgaard, M.4
  • 49
    • 0000243829 scopus 로고
    • PROCHECK: A program to check the stereochemical quality of protein structures
    • Laskowski, R.A., McArthur, M.W., Moss, D.S. & Thornton, J.M. PROCHECK: a program to check the stereochemical quality of protein structures. J. Appl. Crystallogr. 26, 282-291 (1993).
    • (1993) J. Appl. Crystallogr. , vol.26 , pp. 282-291
    • Laskowski, R.A.1    McArthur, M.W.2    Moss, D.S.3    Thornton, J.M.4


* 이 정보는 Elsevier사의 SCOPUS DB에서 KISTI가 분석하여 추출한 것입니다.