메뉴 건너뛰기




Volumn 274, Issue 9, 2007, Pages 2411-2423

Two conserved regions within the tissue-type plasminogen activator gene promoter mediate regulation by brain-derived neurotrophic factor

Author keywords

Brain derived neurotrophic factor; Cortical neurons; Promoter; Tissue type plasminogen activator; Transcription

Indexed keywords

BRAIN DERIVED NEUROTROPHIC FACTOR; BRAIN DERIVED NEUROTROPHIC FACTOR RECEPTOR; TISSUE PLASMINOGEN ACTIVATOR;

EID: 34247339259     PISSN: 1742464X     EISSN: 17424658     Source Type: Journal    
DOI: 10.1111/j.1742-4658.2007.05777.x     Document Type: Article
Times cited : (10)

References (52)
  • 1
    • 16444370452 scopus 로고    scopus 로고
    • Distribution of sympathetic tissue plasminogen activator (tPA) to a distant microvasculature
    • O'Rourke J, Jiang X, Hao Z, Cone RE Hand AR (2005) Distribution of sympathetic tissue plasminogen activator (tPA) to a distant microvasculature. J Neurosci Res 79, 727 733.
    • (2005) J Neurosci Res , vol.79 , pp. 727-733
    • O'Rourke, J.1    Jiang, X.2    Hao, Z.3    Cone, R.E.4    Hand, A.R.5
  • 2
    • 33745584644 scopus 로고    scopus 로고
    • New transgenic evidence for a system of sympathetic axons able to express tissue plasminogen activator (t-PA) within arterial/arteriolar walls
    • Hao Z, Guo C, Jiang X, Krueger S, Pietri T, Dufour S, Cone RE O'Rourke J (2006) New transgenic evidence for a system of sympathetic axons able to express tissue plasminogen activator (t-PA) within arterial/arteriolar walls. Blood 108, 200 202.
    • (2006) Blood , vol.108 , pp. 200-202
    • Hao, Z.1    Guo, C.2    Jiang, X.3    Krueger, S.4    Pietri, T.5    Dufour, S.6    Cone, R.E.7    O'Rourke, J.8
  • 3
    • 0031038431 scopus 로고    scopus 로고
    • Tissue plasminogen activator (t-PA) is targeted to the regulated secretory pathway. Catecholamine storage vesicles as a reservoir for the rapid release of t-PA
    • Parmer RJ, Mahata M, Mahata S, Sebald MT, O'Connor DT Miles LA (1997) Tissue plasminogen activator (t-PA) is targeted to the regulated secretory pathway. Catecholamine storage vesicles as a reservoir for the rapid release of t-PA. J Biol Chem 272, 1976 1982.
    • (1997) J Biol Chem , vol.272 , pp. 1976-1982
    • Parmer, R.J.1    Mahata, M.2    Mahata, S.3    Sebald, M.T.4    O'Connor, D.T.5    Miles, L.A.6
  • 4
    • 0024267754 scopus 로고
    • Tissue plasminogen activator binding to mouse cerebellar granule neurons
    • Verrall S Seeds NW (1988) Tissue plasminogen activator binding to mouse cerebellar granule neurons. J Neurosci Res 21, 420 425.
    • (1988) J Neurosci Res , vol.21 , pp. 420-425
    • Verrall, S.1    Seeds, N.W.2
  • 5
    • 0027535364 scopus 로고
    • Tissue-plasminogen activator is induced as an immediate-early gene during seizure, kindling and long-term potentiation
    • Qian Z, Gilbert ME, Colicos MA, Kandel ER Kuhl D (1993) Tissue-plasminogen activator is induced as an immediate-early gene during seizure, kindling and long-term potentiation. Nature 361, 453 457.
    • (1993) Nature , vol.361 , pp. 453-457
    • Qian, Z.1    Gilbert, M.E.2    Colicos, M.A.3    Kandel, E.R.4    Kuhl, D.5
  • 6
    • 0034064883 scopus 로고    scopus 로고
    • Delayed and blunted induction of mRNA for tissue plasminogen activator in the brain of old rats following pentylenetetrazole-induced seizure activity
    • Popa-Wagner A, Fischer B, Platt D, Schmoll H Kessler C (2000) Delayed and blunted induction of mRNA for tissue plasminogen activator in the brain of old rats following pentylenetetrazole-induced seizure activity. J Gerontol A Biol Sci Med Sci 55, B242 B248.
    • (2000) J Gerontol a Biol Sci Med Sci , vol.55
    • Popa-Wagner, A.1    Fischer, B.2    Platt, D.3    Schmoll, H.4    Kessler, C.5
  • 7
    • 0029556177 scopus 로고
    • Tissue plasminogen activator induction in Purkinje neurons after cerebellar motor learning
    • Seeds NW, Williams BL Bickford PC (1995) Tissue plasminogen activator induction in Purkinje neurons after cerebellar motor learning. Science 270, 1992 1994.
    • (1995) Science , vol.270 , pp. 1992-1994
    • Seeds, N.W.1    Williams, B.L.2    Bickford, P.C.3
  • 10
    • 0029924091 scopus 로고    scopus 로고
    • A different form of long-lasting potentiation revealed in tissue plasminogen activator mutant mice
    • Frey U, Muller M Kuhl D (1996) A different form of long-lasting potentiation revealed in tissue plasminogen activator mutant mice. J Neurosci 16, 2057 2063.
    • (1996) J Neurosci , vol.16 , pp. 2057-2063
    • Frey, U.1    Muller, M.2    Kuhl, D.3
  • 11
    • 9444240943 scopus 로고    scopus 로고
    • Mice lacking the gene encoding tissue-type plasminogen activator show a selective interference with late-phase long-term potentiation in both Schaffer collateral and mossy fiber pathways
    • Huang YY, Bach ME, Lipp HP, Zhuo M, Wolfer DP, Hawkins RD, Schoonjans L, Kandel ER, Godfraind JM, Mulligan R et al. (1996) Mice lacking the gene encoding tissue-type plasminogen activator show a selective interference with late-phase long-term potentiation in both Schaffer collateral and mossy fiber pathways. Proc Natl Acad Sci USA 93, 8699 8704.
    • (1996) Proc Natl Acad Sci USA , vol.93 , pp. 8699-8704
    • Huang, Y.Y.1    Bach, M.E.2    Lipp, H.P.3    Zhuo, M.4    Wolfer, D.P.5    Hawkins, R.D.6    Schoonjans, L.7    Kandel, E.R.8    Godfraind, J.M.9    Mulligan, R.10
  • 13
    • 0032192461 scopus 로고    scopus 로고
    • Tissue plasminogen activator contributes to the late phase of LTP and to synaptic growth in the hippocampal mossy fiber pathway
    • Baranes D, Lederfein D, Huang YY, Chen M, Bailey CH Kandel ER (1998) Tissue plasminogen activator contributes to the late phase of LTP and to synaptic growth in the hippocampal mossy fiber pathway. Neuron 21, 813 825.
    • (1998) Neuron , vol.21 , pp. 813-825
    • Baranes, D.1    Lederfein, D.2    Huang, Y.Y.3    Chen, M.4    Bailey, C.H.5    Kandel, E.R.6
  • 14
    • 0035829385 scopus 로고    scopus 로고
    • Evidence for impairment of behavioural inhibition in performance of operant tasks in tPA-/- mice
    • Ripley TL, Horwood JM Stephens DN (2001) Evidence for impairment of behavioural inhibition in performance of operant tasks in tPA-/- mice. Behav Brain Res 125, 215 227.
    • (2001) Behav Brain Res , vol.125 , pp. 215-227
    • Ripley, T.L.1    Horwood, J.M.2    Stephens, D.N.3
  • 15
    • 0037056011 scopus 로고    scopus 로고
    • Rapid, specific and active site-catalyzed effect of tissue-plasminogen activator on hippocampus-dependent learning in mice
    • Pawlak R, Nagai N, Urano T, Napiorkowska-Pawlak D, Ihara H, Takada Y, Collen D Takada A (2002) Rapid, specific and active site-catalyzed effect of tissue-plasminogen activator on hippocampus-dependent learning in mice. Neuroscience 113, 995 1001.
    • (2002) Neuroscience , vol.113 , pp. 995-1001
    • Pawlak, R.1    Nagai, N.2    Urano, T.3    Napiorkowska-Pawlak, D.4    Ihara, H.5    Takada, Y.6    Collen, D.7    Takada, A.8
  • 16
    • 0042237885 scopus 로고    scopus 로고
    • Absence of tissue plasminogen activator gene or activity impairs mouse cerebellar motor learning
    • Seeds NW, Basham ME Ferguson JE (2003) Absence of tissue plasminogen activator gene or activity impairs mouse cerebellar motor learning. J Neurosci 23, 7368 7375.
    • (2003) J Neurosci , vol.23 , pp. 7368-7375
    • Seeds, N.W.1    Basham, M.E.2    Ferguson, J.E.3
  • 17
  • 18
    • 4444379260 scopus 로고    scopus 로고
    • Tissue plasminogen activator and neuroserpin are widely expressed in the human central nervous system
    • Teesalu T, Kulla A, Simisker A, Siren V, Lawrence DA, Asser T Vaheri A (2004) Tissue plasminogen activator and neuroserpin are widely expressed in the human central nervous system. Thromb Haemostat 92, 358 368.
    • (2004) Thromb Haemostat , vol.92 , pp. 358-368
    • Teesalu, T.1    Kulla, A.2    Simisker, A.3    Siren, V.4    Lawrence, D.A.5    Asser, T.6    Vaheri, A.7
  • 19
    • 0032906124 scopus 로고    scopus 로고
    • BDNF stimulates expression, activity and release of tissue-type plasminogen activator in mouse cortical neurons
    • Fiumelli H, Jabaudon D, Magistretti PJ Martin JL (1999) BDNF stimulates expression, activity and release of tissue-type plasminogen activator in mouse cortical neurons. Eur J Neurosci 11, 1639 1646.
    • (1999) Eur J Neurosci , vol.11 , pp. 1639-1646
    • Fiumelli, H.1    Jabaudon, D.2    Magistretti, P.J.3    Martin, J.L.4
  • 20
    • 8444234659 scopus 로고    scopus 로고
    • Regulation of late-phase LTP and long-term memory in normal and aging hippocampus: Role of secreted proteins tPA and BDNF
    • Pang PT Lu B (2004) Regulation of late-phase LTP and long-term memory in normal and aging hippocampus: role of secreted proteins tPA and BDNF. Ageing Res Rev 3, 407 430.
    • (2004) Ageing Res Rev , vol.3 , pp. 407-430
    • Pang, P.T.1    Lu, B.2
  • 21
    • 25644460530 scopus 로고    scopus 로고
    • Gene expression profiling of facilitated L-LTP in VP16-CREB mice reveals that BDNF is critical for the maintenance of LTP and its synaptic capture
    • Barco A, Patterson S, Alarcon JM, Gromova P, Mata-Roig M, Morozov A Kandel ER (2005) Gene expression profiling of facilitated L-LTP in VP16-CREB mice reveals that BDNF is critical for the maintenance of LTP and its synaptic capture. Neuron 48, 123 137.
    • (2005) Neuron , vol.48 , pp. 123-137
    • Barco, A.1    Patterson, S.2    Alarcon, J.M.3    Gromova, P.4    Mata-Roig, M.5    Morozov, A.6    Kandel, E.R.7
  • 23
    • 0025072939 scopus 로고
    • The structure of the TATA-less rat tissue-type plasminogen activator gene. Species-specific sequence divergences in the promoter predict differences in regulation of gene expression
    • Feng P, Ohlsson M Ny T (1990) The structure of the TATA-less rat tissue-type plasminogen activator gene. Species-specific sequence divergences in the promoter predict differences in regulation of gene expression. J Biol Chem 265, 2022 2027.
    • (1990) J Biol Chem , vol.265 , pp. 2022-2027
    • Feng, P.1    Ohlsson, M.2    Ny, T.3
  • 26
    • 0028235134 scopus 로고
    • A novel regulatory sequence affecting the constitutive expression of tissue plasminogen activator (tPA) gene in human melanoma (Bowes) cells
    • Fujiwara J, Kimura T, Ayusawa D Oishi M (1994) A novel regulatory sequence affecting the constitutive expression of tissue plasminogen activator (tPA) gene in human melanoma (Bowes) cells. J Biol Chem 269, 18558 18562.
    • (1994) J Biol Chem , vol.269 , pp. 18558-18562
    • Fujiwara, J.1    Kimura, T.2    Ayusawa, D.3    Oishi, M.4
  • 27
    • 24144483438 scopus 로고    scopus 로고
    • BDNF function in adult synaptic plasticity: The synaptic consolidation hypothesis
    • Bramham CR Messaoudi E (2005) BDNF function in adult synaptic plasticity: the synaptic consolidation hypothesis. Prog Neurobiol 76, 99 125.
    • (2005) Prog Neurobiol , vol.76 , pp. 99-125
    • Bramham, C.R.1    Messaoudi, E.2
  • 28
    • 33746080943 scopus 로고    scopus 로고
    • Tissue-type plasminogen activator: A multifaceted modulator of neurotransmission and synaptic plasticity
    • Samson AL Medcalf RL (2006) Tissue-type plasminogen activator: a multifaceted modulator of neurotransmission and synaptic plasticity. Neuron 50, 673 678.
    • (2006) Neuron , vol.50 , pp. 673-678
    • Samson, A.L.1    Medcalf, R.L.2
  • 29
    • 0029934148 scopus 로고    scopus 로고
    • Differential binding of cAMP-responsive-element (CRE)-binding protein-1 and activating transcription factor-2 to a CRE-like element in the human tissue-type plasminogen activator (t-PA) gene promoter correlates with opposite regulation of t-PA by phorbol ester in HT-1080 and HeLa cells
    • Costa M Medcalf RL (1996) Differential binding of cAMP-responsive-element (CRE)-binding protein-1 and activating transcription factor-2 to a CRE-like element in the human tissue-type plasminogen activator (t-PA) gene promoter correlates with opposite regulation of t-PA by phorbol ester in HT-1080 and HeLa cells. Eur J Biochem 237, 532 538.
    • (1996) Eur J Biochem , vol.237 , pp. 532-538
    • Costa, M.1    Medcalf, R.L.2
  • 30
    • 23844490237 scopus 로고    scopus 로고
    • In vivo and in vitro analysis of the human tissue-type plasminogen activator gene promoter in neuroblastomal cell lines: Evidence for a functional upstream kappaB element
    • Lux W, Klobeck HG, Daniel PB, Costa M, Medcalf RL Schleuning WD (2005) In vivo and in vitro analysis of the human tissue-type plasminogen activator gene promoter in neuroblastomal cell lines: evidence for a functional upstream kappaB element. J Thromb Haemostat 3, 1009 1017.
    • (2005) J Thromb Haemostat , vol.3 , pp. 1009-1017
    • Lux, W.1    Klobeck, H.G.2    Daniel, P.B.3    Costa, M.4    Medcalf, R.L.5    Schleuning, W.D.6
  • 31
    • 0037899263 scopus 로고    scopus 로고
    • N-Methyl-d-aspartate and TrkB receptor activation in cerebellar granule cells: An in vitro model of preconditioning to stimulate intrinsic survival pathways in neurons
    • [discussion 159-160].
    • Jiang X, Zhu D, Okagaki P, Lipsky R, Wu X, Banaudha K, Mearow K, Strauss KI Marini AM (2003) N-Methyl-d-aspartate and TrkB receptor activation in cerebellar granule cells: an in vitro model of preconditioning to stimulate intrinsic survival pathways in neurons. Ann NY Acad Sci 993, 134 145 [discussion 159-160].
    • (2003) Ann NY Acad Sci , vol.993 , pp. 134-145
    • Jiang, X.1    Zhu, D.2    Okagaki, P.3    Lipsky, R.4    Wu, X.5    Banaudha, K.6    Mearow, K.7    Strauss, K.I.8    Marini, A.M.9
  • 32
    • 0029895777 scopus 로고    scopus 로고
    • Brain-derived neurotrophic factor stimulates AP-1 and cyclic AMP-responsive element dependent transcriptional activity in central nervous system neurons
    • Gaiddon C, Loeffler JP Larmet Y (1996) Brain-derived neurotrophic factor stimulates AP-1 and cyclic AMP-responsive element dependent transcriptional activity in central nervous system neurons. J Neurochem 66, 2279 2286.
    • (1996) J Neurochem , vol.66 , pp. 2279-2286
    • Gaiddon, C.1    Loeffler, J.P.2    Larmet, Y.3
  • 34
    • 17144404150 scopus 로고    scopus 로고
    • Activation of STAT5-dependent transcription by the neurotrophin receptor Trk
    • Klein M, Hempstead BL Teng KK (2005) Activation of STAT5-dependent transcription by the neurotrophin receptor Trk. J Neurobiol 63, 159 171.
    • (2005) J Neurobiol , vol.63 , pp. 159-171
    • Klein, M.1    Hempstead, B.L.2    Teng, K.K.3
  • 35
    • 0041835836 scopus 로고    scopus 로고
    • Brain-derived neurotrophic factor activation of NFAT (nuclear factor of activated T-cells)-dependent transcription: A role for the transcription factor NFATc4 in neurotrophin-mediated gene expression
    • Groth RD Mermelstein PG (2003) Brain-derived neurotrophic factor activation of NFAT (nuclear factor of activated T-cells)-dependent transcription: a role for the transcription factor NFATc4 in neurotrophin-mediated gene expression. J Neurosci 23, 8125 8134.
    • (2003) J Neurosci , vol.23 , pp. 8125-8134
    • Groth, R.D.1    Mermelstein, P.G.2
  • 36
    • 30344442102 scopus 로고    scopus 로고
    • CCAAT/enhancer binding proteins and interferon signaling pathways
    • Kalvakolanu DV Roy SK (2005) CCAAT/enhancer binding proteins and interferon signaling pathways. J Interferon Cytokine Res 25, 757 769.
    • (2005) J Interferon Cytokine Res , vol.25 , pp. 757-769
    • Kalvakolanu, D.V.1    Roy, S.K.2
  • 37
    • 0347298798 scopus 로고    scopus 로고
    • Identification of three NFAT binding motifs in the 5′-upstream region of the human CD3gamma gene that differentially bind NFATc1, NFATc2, and NF-kappa B p50
    • Badran BM, Wolinsky SM, Burny A Willard-Gallo KE (2002) Identification of three NFAT binding motifs in the 5′-upstream region of the human CD3gamma gene that differentially bind NFATc1, NFATc2, and NF-kappa B p50. J Biol Chem 277, 47136 47148.
    • (2002) J Biol Chem , vol.277 , pp. 47136-47148
    • Badran, B.M.1    Wolinsky, S.M.2    Burny, A.3    Willard-Gallo, K.E.4
  • 38
    • 0036421256 scopus 로고    scopus 로고
    • Control elements between -9.5 and -3.0 kb in the human tissue-type plasminogen activator gene promoter direct spatial and inducible expression to the murine brain
    • Yu H, Schleuning WD, Michl M, Liberatore G, Tan SS Medcalf RL (2001) Control elements between -9.5 and -3.0 kb in the human tissue-type plasminogen activator gene promoter direct spatial and inducible expression to the murine brain. Eur J Neurosci 14, 799 808.
    • (2001) Eur J Neurosci , vol.14 , pp. 799-808
    • Yu, H.1    Schleuning, W.D.2    Michl, M.3    Liberatore, G.4    Tan, S.S.5    Medcalf, R.L.6
  • 40
    • 0028043836 scopus 로고
    • The mouse tissue plasminogen activator gene 5′ flanking region directs appropriate expression in development and a seizure-enhanced response in the CNS
    • Carroll PM, Tsirka SE, Richards WG, Frohman MA Strickland S (1994) The mouse tissue plasminogen activator gene 5′ flanking region directs appropriate expression in development and a seizure-enhanced response in the CNS. Development 120, 3173 3183.
    • (1994) Development , vol.120 , pp. 3173-3183
    • Carroll, P.M.1    Tsirka, S.E.2    Richards, W.G.3    Frohman, M.A.4    Strickland, S.5
  • 41
    • 0347694856 scopus 로고    scopus 로고
    • Neurotrophins elevate cAMP to reach a threshold required to overcome inhibition by MAG through extracellular signal-regulated kinase-dependent inhibition of phosphodiesterase
    • Gao Y, Nikulina E, Mellado W Filbin MT (2003) Neurotrophins elevate cAMP to reach a threshold required to overcome inhibition by MAG through extracellular signal-regulated kinase-dependent inhibition of phosphodiesterase. J Neurosci 23, 11770 11777.
    • (2003) J Neurosci , vol.23 , pp. 11770-11777
    • Gao, Y.1    Nikulina, E.2    Mellado, W.3    Filbin, M.T.4
  • 42
    • 0035813199 scopus 로고    scopus 로고
    • Protein synthesis-dependent and -independent regulation of hippocampal synapses by brain-derived neurotrophic factor
    • Tartaglia NJ, Tyler WJ, Neale E, Pozzo-Miller L Lu B (2001) Protein synthesis-dependent and -independent regulation of hippocampal synapses by brain-derived neurotrophic factor. J Biol Chem 276, 37585 37593.
    • (2001) J Biol Chem , vol.276 , pp. 37585-37593
    • Tartaglia, N.J.1    Tyler, W.J.2    Neale, E.3    Pozzo-Miller, L.4    Lu, B.5
  • 43
    • 13244279632 scopus 로고    scopus 로고
    • Cyclic AMP controls BDNF-induced TrkB phosphorylation and dendritic spine formation in mature hippocampal neurons
    • Ji Y, Pang PT, Feng L Lu B (2005) Cyclic AMP controls BDNF-induced TrkB phosphorylation and dendritic spine formation in mature hippocampal neurons. Nat Neurosci 8, 164 172.
    • (2005) Nat Neurosci , vol.8 , pp. 164-172
    • Ji, Y.1    Pang, P.T.2    Feng, L.3    Lu, B.4
  • 44
    • 0035798238 scopus 로고    scopus 로고
    • The molecular biology of memory storage: A dialogue between genes and synapses
    • Kandel ER (2001) The molecular biology of memory storage: a dialogue between genes and synapses. Science 294, 1030 1038.
    • (2001) Science , vol.294 , pp. 1030-1038
    • Kandel, E.R.1
  • 45
    • 0034897668 scopus 로고    scopus 로고
    • Synergistic induction of t-PA by vascular endothelial growth factor and basic fibroblast growth factor and localization of t-PA to Weibel-Palade bodies in bovine microvascular endothelial cells
    • Pepper MS, Rosnoblet C, Di Sanza C Kruithof EK (2001) Synergistic induction of t-PA by vascular endothelial growth factor and basic fibroblast growth factor and localization of t-PA to Weibel-Palade bodies in bovine microvascular endothelial cells. Thromb Haemostat 86, 702 709.
    • (2001) Thromb Haemostat , vol.86 , pp. 702-709
    • Pepper, M.S.1    Rosnoblet, C.2    Di Sanza, C.3    Kruithof, E.K.4
  • 47
    • 0037314824 scopus 로고    scopus 로고
    • Tissue plasminogen activator in the amygdala is critical for stress-induced anxiety-like behavior
    • Pawlak R, Magarinos AM, Melchor J, McEwen B Strickland S (2003) Tissue plasminogen activator in the amygdala is critical for stress-induced anxiety-like behavior. Nat Neurosci 6, 168 174.
    • (2003) Nat Neurosci , vol.6 , pp. 168-174
    • Pawlak, R.1    Magarinos, A.M.2    Melchor, J.3    McEwen, B.4    Strickland, S.5
  • 48
    • 0034729210 scopus 로고    scopus 로고
    • Tissue plasminogen activator-mediated fibrinolysis protects against axonal degeneration and demyelination after sciatic nerve injury
    • Akassoglou K, Kombrinck KW, Degen JL Strickland S (2000) Tissue plasminogen activator-mediated fibrinolysis protects against axonal degeneration and demyelination after sciatic nerve injury. J Cell Biol 149, 1157 1166.
    • (2000) J Cell Biol , vol.149 , pp. 1157-1166
    • Akassoglou, K.1    Kombrinck, K.W.2    Degen, J.L.3    Strickland, S.4
  • 49
    • 25144499286 scopus 로고    scopus 로고
    • A role for the plasminogen activator system in inflammation and neurodegeneration in the central nervous system during experimental allergic encephalomyelitis
    • East E, Baker D, Pryce G, Lijnen HR, Cuzner ML Gveric D (2005) A role for the plasminogen activator system in inflammation and neurodegeneration in the central nervous system during experimental allergic encephalomyelitis. Am J Pathol 167, 545 554.
    • (2005) Am J Pathol , vol.167 , pp. 545-554
    • East, E.1    Baker, D.2    Pryce, G.3    Lijnen, H.R.4    Cuzner, M.L.5    Gveric, D.6


* 이 정보는 Elsevier사의 SCOPUS DB에서 KISTI가 분석하여 추출한 것입니다.