메뉴 건너뛰기




Volumn 291, Issue 15, 2016, Pages 8214-8230

Insights into the effects of complement factor H on the assembly and decay of the alternative pathway C3 proconvertase and C3 convertase

Author keywords

[No Author keywords available]

Indexed keywords

BINS; DISSOCIATION;

EID: 84964626186     PISSN: 00219258     EISSN: 1083351X     Source Type: Journal    
DOI: 10.1074/jbc.M115.693119     Document Type: Article
Times cited : (11)

References (61)
  • 1
    • 0035810399 scopus 로고    scopus 로고
    • Complement. First of two parts
    • Walport, M. J. (2001) Complement. First of two parts. N. Engl. J. Med. 344, 1058-1066
    • (2001) N. Engl. J. Med. , vol.344 , pp. 1058-1066
    • Walport, M.J.1
  • 2
    • 0035849176 scopus 로고    scopus 로고
    • Complement. Second of two parts
    • Walport, M. J. (2001) Complement. Second of two parts. N. Engl. J. Med. 344, 1140-1144
    • (2001) N. Engl. J. Med. , vol.344 , pp. 1140-1144
    • Walport, M.J.1
  • 3
    • 77955883153 scopus 로고    scopus 로고
    • Complement: A key system for immune surveillance and homeostasis
    • Ricklin, D., Hajishengallis, G., Yang, K., and Lambris, J. D. (2010) Complement: a key system for immune surveillance and homeostasis. Nat. Immunol. 11, 785-797
    • (2010) Nat. Immunol. , vol.11 , pp. 785-797
    • Ricklin, D.1    Hajishengallis, G.2    Yang, K.3    Lambris, J.D.4
  • 4
    • 70350279315 scopus 로고    scopus 로고
    • Atypical hemolytic-uremic syndrome
    • Noris, M., and Remuzzi, G. (2009) Atypical hemolytic-uremic syndrome. N. Engl. J. Med. 361, 1676-1687
    • (2009) N. Engl. J. Med. , vol.361 , pp. 1676-1687
    • Noris, M.1    Remuzzi, G.2
  • 5
    • 84886309053 scopus 로고    scopus 로고
    • Overview of complement activation and regulation
    • Noris, M., and Remuzzi, G. (2013) Overview of complement activation and regulation. Semin. Nephrol. 33, 479-492
    • (2013) Semin. Nephrol. , vol.33 , pp. 479-492
    • Noris, M.1    Remuzzi, G.2
  • 6
    • 78649450284 scopus 로고    scopus 로고
    • Lessons from functional and structural analyses of disease-associated genetic variants in the complement alternative pathway
    • Rodríguez de Córdoba, S., Harris, C. L., Morgan, B. P., and Llorca, O. (2011) Lessons from functional and structural analyses of disease-associated genetic variants in the complement alternative pathway. Biochim. Biophys. Acta 1812, 12-22
    • (2011) Biochim. Biophys. Acta , vol.1812 , pp. 12-22
    • Rodríguez De Córdoba, S.1    Harris, C.L.2    Morgan, B.P.3    Llorca, O.4
  • 7
    • 84867994785 scopus 로고    scopus 로고
    • Pathogenesis of the C3 glomerulopathies and reclassification of MPGN
    • Bomback, A. S., and Appel, G. B. (2012) Pathogenesis of the C3 glomerulopathies and reclassification of MPGN. Nat. Rev. Nephrol. 8, 634-642
    • (2012) Nat. Rev. Nephrol. , vol.8 , pp. 634-642
    • Bomback, A.S.1    Appel, G.B.2
  • 8
    • 84864564017 scopus 로고    scopus 로고
    • C3 glomerulonephritis: Clinicopathological findings, complement abnormalities, glomerular proteomic profile, treatment, and follow-up
    • Sethi, S., Fervenza, F. C., Zhang, Y., Zand, L., Vrana, J. A., Nasr, S. H., Theis, J. D., Dogan, A., and Smith, R. J. (2012) C3 glomerulonephritis: clinicopathological findings, complement abnormalities, glomerular proteomic profile, treatment, and follow-up. Kidney Int. 82, 465-473
    • (2012) Kidney Int. , vol.82 , pp. 465-473
    • Sethi, S.1    Fervenza, F.C.2    Zhang, Y.3    Zand, L.4    Vrana, J.A.5    Nasr, S.H.6    Theis, J.D.7    Dogan, A.8    Smith, R.J.9
  • 9
    • 84936985072 scopus 로고    scopus 로고
    • Age-related macular degeneration and the role of the complement system
    • McHarg, S., Clark, S. J., Day, A. J., and Bishop, P. N. (2015) Age-related macular degeneration and the role of the complement system. Mol. Immunol. 67, 43-50
    • (2015) Mol. Immunol. , vol.67 , pp. 43-50
    • McHarg, S.1    Clark, S.J.2    Day, A.J.3    Bishop, P.N.4
  • 10
    • 59449088846 scopus 로고    scopus 로고
    • Eculizumab for congenital atypical hemolytic-uremic syndrome
    • Gruppo, R. A., and Rother, R. P. (2009) Eculizumab for congenital atypical hemolytic-uremic syndrome. N. Engl. J. Med. 360, 544-546
    • (2009) N. Engl. J. Med. , vol.360 , pp. 544-546
    • Gruppo, R.A.1    Rother, R.P.2
  • 12
    • 84867993256 scopus 로고    scopus 로고
    • Use of eculizumab for atypical haemolytic uraemic syndrome and C3 glomerulopathies
    • Zuber, J., Fakhouri, F., Roumenina, L. T., Loirat, C., Frémeaux-Bacchi, V., French Study Group for aHUS/C3G. (2012) Use of eculizumab for atypical haemolytic uraemic syndrome and C3 glomerulopathies. Nat. Rev. Nephrol. 8, 643-657
    • (2012) Nat. Rev. Nephrol. , vol.8 , pp. 643-657
    • Zuber, J.1    Fakhouri, F.2    Roumenina, L.T.3    Loirat, C.4    Frémeaux-Bacchi, V.5
  • 13
    • 84858671433 scopus 로고    scopus 로고
    • Eculizumab in a patient with dense-deposit disease
    • Daina, E., Noris, M., and Remuzzi, G. (2012) Eculizumab in a patient with dense-deposit disease. N. Engl. J. Med. 366, 1161-1163
    • (2012) N. Engl. J. Med. , vol.366 , pp. 1161-1163
    • Daina, E.1    Noris, M.2    Remuzzi, G.3
  • 14
    • 84902254536 scopus 로고    scopus 로고
    • Treatment of C3 glomerulopathy with complement blockers
    • Vivarelli, M., and Emma, F. (2014) Treatment of C3 glomerulopathy with complement blockers. Semin. Thromb. Hemost. 40, 472-477
    • (2014) Semin. Thromb. Hemost. , vol.40 , pp. 472-477
    • Vivarelli, M.1    Emma, F.2
  • 16
    • 84865549729 scopus 로고    scopus 로고
    • Immunology. The new view of complement
    • Leslie, M. (2012) Immunology. The new view of complement. Science 337, 1034-1037
    • (2012) Science , vol.337 , pp. 1034-1037
    • Leslie, M.1
  • 17
    • 0016703811 scopus 로고
    • Properdin: Binding to C3b and stabilization of the C3b-dependent C3 convertase
    • Fearon, D. T., and Austen, K. F. (1975) Properdin: binding to C3b and stabilization of the C3b-dependent C3 convertase. J. Exp. Med. 142, 856-863
    • (1975) J. Exp. Med. , vol.142 , pp. 856-863
    • Fearon, D.T.1    Austen, K.F.2
  • 18
    • 33644846402 scopus 로고    scopus 로고
    • The role of properdin in the assembly of the alternative pathway C3 convertases of complement
    • Hourcade, D. E. (2006) The role of properdin in the assembly of the alternative pathway C3 convertases of complement. J. Biol. Chem. 281, 2128-2132
    • (2006) J. Biol. Chem. , vol.281 , pp. 2128-2132
    • Hourcade, D.E.1
  • 19
    • 70349437186 scopus 로고    scopus 로고
    • Complement regulators and inhibitory proteins
    • Zipfel, P. F., and Skerka, C. (2009) Complement regulators and inhibitory proteins. Nat. Rev. Immunol. 9, 729-740
    • (2009) Nat. Rev. Immunol. , vol.9 , pp. 729-740
    • Zipfel, P.F.1    Skerka, C.2
  • 20
    • 0017704496 scopus 로고
    • Human complement C3b inactivator: Isolation, characterization, and demonstration of an absolute requirement for the serum protein β1H for cleavage of C3b and C4b in solution
    • Pangburn, M. K., Schreiber, R. D., and Müller-Eberhard, H. J. (1977) Human complement C3b inactivator: isolation, characterization, and demonstration of an absolute requirement for the serum protein β1H for cleavage of C3b and C4b in solution. J. Exp. Med. 146, 257-270
    • (1977) J. Exp. Med. , vol.146 , pp. 257-270
    • Pangburn, M.K.1    Schreiber, R.D.2    Müller-Eberhard, H.J.3
  • 21
    • 0012042656 scopus 로고
    • Control of the amplification convertase of complement by the plasma protein β1H
    • Weiler, J. M., Daha, M. R., Austen, K. F., and Fearon, D. T. (1976) Control of the amplification convertase of complement by the plasma protein β1H. Proc. Natl. Acad. Sci. U.S.A. 73, 3268-3272
    • (1976) Proc. Natl. Acad. Sci. U.S.A. , vol.73 , pp. 3268-3272
    • Weiler, J.M.1    Daha, M.R.2    Austen, K.F.3    Fearon, D.T.4
  • 22
    • 0017139872 scopus 로고
    • Modulation of the alternative complement pathways by β1H globulin
    • Whaley, K., and Ruddy, S. (1976) Modulation of the alternative complement pathways by β1H globulin. J. Exp. Med. 144, 1147-1163
    • (1976) J. Exp. Med. , vol.144 , pp. 1147-1163
    • Whaley, K.1    Ruddy, S.2
  • 23
    • 0018199513 scopus 로고
    • Interaction of β1H globulin with cell-bound C3b: Quantitative analysis of binding and influence of alternative pathway components on binding
    • Conrad, D. H., Carlo, J. R., and Ruddy, S. (1978) Interaction of β1H globulin with cell-bound C3b: quantitative analysis of binding and influence of alternative pathway components on binding. J. Exp. Med. 147, 1792-1805
    • (1978) J. Exp. Med. , vol.147 , pp. 1792-1805
    • Conrad, D.H.1    Carlo, J.R.2    Ruddy, S.3
  • 24
    • 0011579669 scopus 로고
    • Complement C3 convertase: Cell surface restriction of β1H control and generation of restriction on neuraminidase-treated cells
    • Pangburn, M. K., and Müller-Eberhard, H. J. (1978) Complement C3 convertase: cell surface restriction of β1H control and generation of restriction on neuraminidase-treated cells. Proc. Natl. Acad. Sci. U.S.A. 75, 2416-2420
    • (1978) Proc. Natl. Acad. Sci. U.S.A. , vol.75 , pp. 2416-2420
    • Pangburn, M.K.1    Müller-Eberhard, H.J.2
  • 25
    • 0018317786 scopus 로고
    • Human alternative complement pathway: Membrane-associated sialic acid regulates the competition between B and β1H for cell-bound C3b
    • Kazatchkine, M. D., Fearon, D. T., and Austen, K. F. (1979) Human alternative complement pathway: membrane-associated sialic acid regulates the competition between B and β1H for cell-bound C3b. J. Immunol. 122, 75-81
    • (1979) J. Immunol. , vol.122 , pp. 75-81
    • Kazatchkine, M.D.1    Fearon, D.T.2    Austen, K.F.3
  • 26
    • 69449090499 scopus 로고    scopus 로고
    • The disease-protective complement factor H allotypic variant Ile62 shows increased binding affinity for C3b and enhanced cofactor activity
    • Tortajada, A., Montes, T., Martínez-Barricarte, R., Morgan, B. P., Harris, C. L., and de Córdoba, S. R. (2009) The disease-protective complement factor H allotypic variant Ile62 shows increased binding affinity for C3b and enhanced cofactor activity. Hum. Mol. Genet. 18, 3452-3461
    • (2009) Hum. Mol. Genet. , vol.18 , pp. 3452-3461
    • Tortajada, A.1    Montes, T.2    Martínez-Barricarte, R.3    Morgan, B.P.4    Harris, C.L.5    De Córdoba, S.R.6
  • 27
    • 0032997017 scopus 로고    scopus 로고
    • Decay acceleration of the complement alternative pathway C3 convertase
    • Hourcade, D. E., Mitchell, L. M., and Medof, M. E. (1999) Decay acceleration of the complement alternative pathway C3 convertase. Immunopharmacology 42, 167-173
    • (1999) Immunopharmacology , vol.42 , pp. 167-173
    • Hourcade, D.E.1    Mitchell, L.M.2    Medof, M.E.3
  • 28
    • 84878910014 scopus 로고    scopus 로고
    • Functional anatomy of complement factor H
    • Makou, E., Herbert, A. P., and Barlow, P. N. (2013) Functional anatomy of complement factor H. Biochemistry 52, 3949-3962
    • (2013) Biochemistry , vol.52 , pp. 3949-3962
    • Makou, E.1    Herbert, A.P.2    Barlow, P.N.3
  • 29
    • 36849044745 scopus 로고    scopus 로고
    • Translational mini-review series on complement factor H: Structural and functional correlations for factor H
    • Schmidt, C. Q., Herbert, A. P., Hocking, H. G., Uhrín, D., and Barlow, P. N. (2008) Translational mini-review series on complement factor H: structural and functional correlations for factor H. Clin. Exp. Immunol 151, 14-24
    • (2008) Clin. Exp. Immunol , vol.151 , pp. 14-24
    • Schmidt, C.Q.1    Herbert, A.P.2    Hocking, H.G.3    Uhrín, D.4    Barlow, P.N.5
  • 31
    • 0029086243 scopus 로고
    • Analysis of the short consensus repeats of human complement factor B by site-directed mutagenesis
    • Hourcade, D. E., Wagner, L. M., and Oglesby, T. J. (1995) Analysis of the short consensus repeats of human complement factor B by site-directed mutagenesis. J. Biol. Chem. 270, 19716-19722
    • (1995) J. Biol. Chem. , vol.270 , pp. 19716-19722
    • Hourcade, D.E.1    Wagner, L.M.2    Oglesby, T.J.3
  • 32
    • 0037059776 scopus 로고    scopus 로고
    • Decay-accelerating factor (DAF), complement receptor 1 (CR1), and factor H dissociate the complement AP C3 convertase (C3bBb) via sites on the type A domain of Bb
    • Hourcade, D. E., Mitchell, L., Kuttner-Kondo, L. A., Atkinson, J. P., and Medof, M. E. (2002) Decay-accelerating factor (DAF), complement receptor 1 (CR1), and factor H dissociate the complement AP C3 convertase (C3bBb) via sites on the type A domain of Bb. J. Biol. Chem. 277, 1107-1112
    • (2002) J. Biol. Chem. , vol.277 , pp. 1107-1112
    • Hourcade, D.E.1    Mitchell, L.2    Kuttner-Kondo, L.A.3    Atkinson, J.P.4    Medof, M.E.5
  • 33
    • 80053898466 scopus 로고    scopus 로고
    • Access to the complement factor B scissile bond is facilitated by association of factor B with C3b protein
    • Hourcade, D. E., and Mitchell, L. M. (2011) Access to the complement factor B scissile bond is facilitated by association of factor B with C3b protein. J. Biol. Chem. 286, 35725-35732
    • (2011) J. Biol. Chem. , vol.286 , pp. 35725-35732
    • Hourcade, D.E.1    Mitchell, L.M.2
  • 34
    • 0020438716 scopus 로고
    • C3 convertase of human complement: Enhanced formation and stability of the enzyme generated with nickel instead of magnesium
    • Fishelson, Z., and Müller-Eberhard, H. J. (1982) C3 convertase of human complement: enhanced formation and stability of the enzyme generated with nickel instead of magnesium. J. Immunol. 129, 2603-2607
    • (1982) J. Immunol. , vol.129 , pp. 2603-2607
    • Fishelson, Z.1    Müller-Eberhard, H.J.2
  • 35
    • 84883693955 scopus 로고    scopus 로고
    • Properdin and factor H: Opposing players on the alternative complement pathway "see-saw"
    • Kouser, L., Abdul-Aziz, M., Nayak, A., Stover, C. M., Sim, R. B., and Kishore, U. (2013) Properdin and factor H: opposing players on the alternative complement pathway "see-saw". Front. Immunol. 4, 93
    • (2013) Front. Immunol. , vol.4 , pp. 93
    • Kouser, L.1    Abdul-Aziz, M.2    Nayak, A.3    Stover, C.M.4    Sim, R.B.5    Kishore, U.6
  • 36
    • 0028245945 scopus 로고
    • Selective disappearance of C3NeF IgG autoantibody in the plasma of a patient with membranoproliferative glomerulonephritis following renal transplantation
    • Frémeaux-Bacchi, V., Weiss, L., Brun, P., and Kazatchkine, M. D. (1994) Selective disappearance of C3NeF IgG autoantibody in the plasma of a patient with membranoproliferative glomerulonephritis following renal transplantation. Nephrol. Dial. Transplant. 9, 811-814
    • (1994) Nephrol. Dial. Transplant. , vol.9 , pp. 811-814
    • Frémeaux-Bacchi, V.1    Weiss, L.2    Brun, P.3    Kazatchkine, M.D.4
  • 37
    • 58849109079 scopus 로고    scopus 로고
    • 3D structure of the C3bB complex provides insights into the activation and regulation of the complement alternative pathway convertase
    • Torreira, E., Tortajada, A., Montes, T., Rodríguez de Córdoba, S., and Llorca, O. (2009) 3D structure of the C3bB complex provides insights into the activation and regulation of the complement alternative pathway convertase. Proc. Natl. Acad. Sci. U.S.A. 106, 882-887
    • (2009) Proc. Natl. Acad. Sci. U.S.A. , vol.106 , pp. 882-887
    • Torreira, E.1    Tortajada, A.2    Montes, T.3    Rodríguez De Córdoba, S.4    Llorca, O.5
  • 38
    • 13244291613 scopus 로고    scopus 로고
    • Molecular dissection of interactions between components of the alternative pathway of complement and decay accelerating factor (CD55)
    • Harris, C. L., Abbott, R. J., Smith, R. A., Morgan, B. P., and Lea, S. M. (2005) Molecular dissection of interactions between components of the alternative pathway of complement and decay accelerating factor (CD55). J. Biol. Chem. 280, 2569-2578
    • (2005) J. Biol. Chem. , vol.280 , pp. 2569-2578
    • Harris, C.L.1    Abbott, R.J.2    Smith, R.A.3    Morgan, B.P.4    Lea, S.M.5
  • 40
    • 84882447182 scopus 로고    scopus 로고
    • Structural basis for the stabilization of the complement alternative pathway C3 convertase by properdin
    • Alcorlo, M., Tortajada, A., Rodríguez de Córdoba, S., and Llorca, O. (2013) Structural basis for the stabilization of the complement alternative pathway C3 convertase by properdin. Proc. Natl. Acad. Sci. U.S.A. 110, 13504-13509
    • (2013) Proc. Natl. Acad. Sci. U.S.A. , vol.110 , pp. 13504-13509
    • Alcorlo, M.1    Tortajada, A.2    Rodríguez De Córdoba, S.3    Llorca, O.4
  • 45
    • 0027339666 scopus 로고
    • Conformational epitopes of C3 reflecting its mode of binding to an artificial polymer surface
    • Nilsson, U. R., Storm, K. E., Elwing, H., and Nilsson, B. (1993) Conformational epitopes of C3 reflecting its mode of binding to an artificial polymer surface. Mol. Immunol. 30, 211-219
    • (1993) Mol. Immunol. , vol.30 , pp. 211-219
    • Nilsson, U.R.1    Storm, K.E.2    Elwing, H.3    Nilsson, B.4
  • 50
    • 33646040989 scopus 로고    scopus 로고
    • CRIT peptide interacts with factor B and interferes with alternative pathway activation
    • Hui, K. M., Magnadóttir, B., Schifferli, J. A., and Inal, J. M. (2006) CRIT peptide interacts with factor B and interferes with alternative pathway activation. Biochem. Biophys. Res. Commun. 344, 308-314
    • (2006) Biochem. Biophys. Res. Commun. , vol.344 , pp. 308-314
    • Hui, K.M.1    Magnadóttir, B.2    Schifferli, J.A.3    Inal, J.M.4
  • 52
    • 73349135351 scopus 로고    scopus 로고
    • Coexistence of closed and open conformations of complement factor B in the alternative pathway C3bB(Mg2+) proconvertase
    • Torreira, E., Tortajada, A., Montes, T., Rodríguez de Córdoba, S., and Llorca, O. (2009) Coexistence of closed and open conformations of complement factor B in the alternative pathway C3bB(Mg2+) proconvertase. J. Immunol. 183, 7347-7351
    • (2009) J. Immunol. , vol.183 , pp. 7347-7351
    • Torreira, E.1    Tortajada, A.2    Montes, T.3    Rodríguez De Córdoba, S.4    Llorca, O.5
  • 54
    • 78650638514 scopus 로고    scopus 로고
    • Structures of C3b in complex with factors B andDgive insight into complement convertase formation
    • Forneris, F., Ricklin, D., Wu, J., Tzekou, A., Wallace, R. S., Lambris, J. D., and Gros, P. (2010) Structures of C3b in complex with factors B andDgive insight into complement convertase formation. Science 330, 1816-1820
    • (2010) Science , vol.330 , pp. 1816-1820
    • Forneris, F.1    Ricklin, D.2    Wu, J.3    Tzekou, A.4    Wallace, R.S.5    Lambris, J.D.6    Gros, P.7
  • 56
    • 67649230210 scopus 로고    scopus 로고
    • Structure of complement fragment C3b-factor H and implications for host protection by complement regulators
    • Wu, J., Wu, Y. Q., Ricklin, D., Janssen, B. J., Lambris, J. D., and Gros, P. (2009) Structure of complement fragment C3b-factor H and implications for host protection by complement regulators. Nat. Immunol. 10, 728-733
    • (2009) Nat. Immunol. , vol.10 , pp. 728-733
    • Wu, J.1    Wu, Y.Q.2    Ricklin, D.3    Janssen, B.J.4    Lambris, J.D.5    Gros, P.6
  • 57
    • 0033921990 scopus 로고    scopus 로고
    • Host recognition and target differentiation by factor H, a regulator of the alternative pathway of complement
    • Pangburn, M. K. (2000) Host recognition and target differentiation by factor H, a regulator of the alternative pathway of complement. Immunopharmacology 49, 149-157
    • (2000) Immunopharmacology , vol.49 , pp. 149-157
    • Pangburn, M.K.1
  • 59
    • 67650129379 scopus 로고    scopus 로고
    • Mutations of factorHimpair regulation of surface-bound C3b by three mechanisms in atypical hemolytic uremic syndrome
    • Lehtinen, M. J., Rops, A. L., Isenman, D. E., van der Vlag, J., and Jokiranta, T. S. (2009) Mutations of factorHimpair regulation of surface-bound C3b by three mechanisms in atypical hemolytic uremic syndrome. J. Biol. Chem. 284, 15650-15658
    • (2009) J. Biol. Chem. , vol.284 , pp. 15650-15658
    • Lehtinen, M.J.1    Rops, A.L.2    Isenman, D.E.3    Van Der Vlag, J.4    Jokiranta, T.S.5
  • 60
    • 69249220131 scopus 로고    scopus 로고
    • Insights into complement convertase formation based on the structure of the factor B-cobra venom factor complex
    • Janssen, B. J., Gomes, L., Koning, R. I., Svergun, D. I., Koster, A. J., Fritzinger, D. C., Vogel, C. W., and Gros, P. (2009) Insights into complement convertase formation based on the structure of the factor B-cobra venom factor complex. EMBO J. 28, 2469-2478
    • (2009) EMBO J. , vol.28 , pp. 2469-2478
    • Janssen, B.J.1    Gomes, L.2    Koning, R.I.3    Svergun, D.I.4    Koster, A.J.5    Fritzinger, D.C.6    Vogel, C.W.7    Gros, P.8
  • 61
    • 33750859976 scopus 로고    scopus 로고
    • Structure of C3b reveals conformational changes that underlie complement activity
    • Janssen, B. J., Christodoulidou, A., McCarthy, A., Lambris, J. D., and Gros, P. (2006) Structure of C3b reveals conformational changes that underlie complement activity. Nature 444, 213-216
    • (2006) Nature , vol.444 , pp. 213-216
    • Janssen, B.J.1    Christodoulidou, A.2    McCarthy, A.3    Lambris, J.D.4    Gros, P.5


* 이 정보는 Elsevier사의 SCOPUS DB에서 KISTI가 분석하여 추출한 것입니다.