메뉴 건너뛰기




Volumn 32, Issue 37, 2012, Pages 12854-12861

Integrin dynamics produce a delayed stage of long-term potentiation and memory consolidation

Author keywords

[No Author keywords available]

Indexed keywords

ANISOMYCIN; BETA1 INTEGRIN; BREFELDIN A; FOCAL ADHESION KINASE; FOCAL ADHESION KINASE 2; POSTSYNAPTIC DENSITY PROTEIN 95;

EID: 84866254118     PISSN: 02706474     EISSN: 15292401     Source Type: Journal    
DOI: 10.1523/JNEUROSCI.2024-12.2012     Document Type: Article
Times cited : (77)

References (58)
  • 2
    • 79951775349 scopus 로고    scopus 로고
    • The biochemistry of memory: The 26 year journey of a 'new and specific hypothesis'
    • Baudry M, Bi X, Gall C, Lynch G (2011) The biochemistry of memory: the 26 year journey of a 'new and specific hypothesis'. Neurobiol Learn Mem 95:125-133.
    • (2011) Neurobiol Learn Mem , vol.95 , pp. 125-133
    • Baudry, M.1    Bi, X.2    Gall, C.3    Lynch, G.4
  • 3
    • 18844391463 scopus 로고    scopus 로고
    • Integrin signaling cascades are operational in adult hippocampal synapses and modulate NMDA receptor physiology
    • Bernard-Trifilo JA, Kramár EA, Torp R, Lin CY, Pineda EA, Lynch G, Gall CM (2005) Integrin signaling cascades are operational in adult hippocampal synapses and modulate NMDA receptor physiology. J Neurochem 93:834-849.
    • (2005) J Neurochem , vol.93 , pp. 834-849
    • Bernard-Trifilo, J.A.1    Kramár, E.A.2    Torp, R.3    Lin, C.Y.4    Pineda, E.A.5    Lynch, G.6    Gall, C.M.7
  • 4
    • 84863446250 scopus 로고    scopus 로고
    • Nanoscale analysis of structural synaptic plasticity
    • Bourne JN, Harris KM (2012) Nanoscale analysis of structural synaptic plasticity. Curr Opin Neurobiol 22:372-382.
    • (2012) Curr Opin Neurobiol , vol.22 , pp. 372-382
    • Bourne, J.N.1    Harris, K.M.2
  • 5
    • 34447552405 scopus 로고    scopus 로고
    • In vivo roles for matrix metalloproteinase-9 in mature hippocampal synaptic physiology and plasticity
    • Bozdagi O, Nagy V, Kwei KT, Huntley GW (2007) In vivo roles for matrix metalloproteinase-9 in mature hippocampal synaptic physiology and plasticity. J Neurophysiol 98:334-344.
    • (2007) J Neurophysiol , vol.98 , pp. 334-344
    • Bozdagi, O.1    Nagy, V.2    Kwei, K.T.3    Huntley, G.W.4
  • 6
    • 0037904987 scopus 로고    scopus 로고
    • The integrin-actin connection, an eternal love affair
    • Brakebusch C, Fässler R (2003) The integrin-actin connection, an eternal love affair. EMBO J 22:2324-2333.
    • (2003) EMBO J , vol.22 , pp. 2324-2333
    • Brakebusch, C.1    Fässler, R.2
  • 7
    • 56549084442 scopus 로고    scopus 로고
    • Local protein synthesis, actin dynamics, and LTP consolidation
    • Bramham CR (2008) Local protein synthesis, actin dynamics, and LTP consolidation. Curr Opin Neurobiol 18:524-531.
    • (2008) Curr Opin Neurobiol , vol.18 , pp. 524-531
    • Bramham, C.R.1
  • 8
    • 0031610051 scopus 로고    scopus 로고
    • The spacing effect depends on an encoding deficit, retrieval, and time in working memory: Evidence from oncepresented words
    • Braun K, Rubin DC (1998) The spacing effect depends on an encoding deficit, retrieval, and time in working memory: evidence from oncepresented words. Memory 6:37-65.
    • (1998) Memory , vol.6 , pp. 37-65
    • Braun, K.1    Rubin, D.C.2
  • 9
    • 0042125569 scopus 로고    scopus 로고
    • Integrin requirement for hippocampal synaptic plasticity and spatial memory
    • Chan CS, Weeber EJ, Kurup S, Sweatt JD, Davis RL (2003) Integrin requirement for hippocampal synaptic plasticity and spatial memory. J Neurosci 23:7107-7116.
    • (2003) J Neurosci , vol.23 , pp. 7107-7116
    • Chan, C.S.1    Weeber, E.J.2    Kurup, S.3    Sweatt, J.D.4    Davis, R.L.5
  • 10
    • 30644461034 scopus 로고    scopus 로고
    • Beta 1-integrins are required for hippocampal AMPA receptor-dependent synaptic transmission, synaptic plasticity, and working memory
    • Chan CS, Weeber EJ, Zong L, Fuchs E, Sweatt JD, Davis RL (2006) Beta 1-integrins are required for hippocampal AMPA receptor-dependent synaptic transmission, synaptic plasticity, and working memory. J Neurosci 26:223-232.
    • (2006) J Neurosci , vol.26 , pp. 223-232
    • Chan, C.S.1    Weeber, E.J.2    Zong, L.3    Fuchs, E.4    Sweatt, J.D.5    Davis, R.L.6
  • 12
    • 77952976816 scopus 로고    scopus 로고
    • α8-integrins are required for hippocampal long-term potentiation but not for hippocampal-dependent learning
    • Chan CS, Chen H, Bradley A, Dragatsis I, Rosenmund C, Davis RL (2010) α8-integrins are required for hippocampal long-term potentiation but not for hippocampal-dependent learning. Genes Brain Behav 9:402-410.
    • (2010) Genes Brain Behav , vol.9 , pp. 402-410
    • Chan, C.S.1    Chen, H.2    Bradley, A.3    Dragatsis, I.4    Rosenmund, C.5    Davis, R.L.6
  • 13
    • 34249055892 scopus 로고    scopus 로고
    • Changes in synaptic morphology accompany actin signaling during LTP
    • Chen LY, Rex CS, Casale MS, Gall CM, Lynch G (2007) Changes in synaptic morphology accompany actin signaling during LTP. J Neurosci 27:5363-5372.
    • (2007) J Neurosci , vol.27 , pp. 5363-5372
    • Chen, L.Y.1    Rex, C.S.2    Casale, M.S.3    Gall, C.M.4    Lynch, G.5
  • 14
    • 78149482421 scopus 로고    scopus 로고
    • BDNF signaling during learning is regionally differentiated within hippocampus
    • Chen LY, Rex CS, Pham DT, Lynch G, Gall CM (2010a) BDNF signaling during learning is regionally differentiated within hippocampus. J Neurosci 30:15097-15101.
    • (2010) J Neurosci , vol.30 , pp. 15097-15101
    • Chen, L.Y.1    Rex, C.S.2    Pham, D.T.3    Lynch, G.4    Gall, C.M.5
  • 16
    • 77955914539 scopus 로고    scopus 로고
    • Physiological activation of synaptic Rac>PAK (p-21 activated kinase) signaling is defective in a mouse model of fragile X syndrome
    • Chen LY, Rex CS, Babayan AH, Kramár EA, Lynch G, Gall CM, Lauterborn JC (2010c) Physiological activation of synaptic Rac>PAK (p-21 activated kinase) signaling is defective in a mouse model of fragile X syndrome. J Neurosci 30:10977-10984.
    • (2010) J Neurosci , vol.30 , pp. 10977-10984
    • Chen, L.Y.1    Rex, C.S.2    Babayan, A.H.3    Kramár, E.A.4    Lynch, G.5    Gall, C.M.6    Lauterborn, J.C.7
  • 17
    • 0035934293 scopus 로고    scopus 로고
    • Evidence that integrins contribute to multiple stages in the consolidation of long term potentiation in rat hippocampus
    • Chun D, Gall CM, Bi X, Lynch G (2001) Evidence that integrins contribute to multiple stages in the consolidation of long term potentiation in rat hippocampus. Neuroscience 105:815-829.
    • (2001) Neuroscience , vol.105 , pp. 815-829
    • Chun, D.1    Gall, C.M.2    Bi, X.3    Lynch, G.4
  • 18
    • 0025013459 scopus 로고
    • Development of hippocampal long-term potentiation is reduced by recently introduced calpain inhibitors
    • del Cerro S, Larson J, Oliver MW, Lynch G (1990) Development of hippocampal long-term potentiation is reduced by recently introduced calpain inhibitors. Brain Res 530:91-95.
    • (1990) Brain Res , vol.530 , pp. 91-95
    • del Cerro, S.1    Larson, J.2    Oliver, M.W.3    Lynch, G.4
  • 19
    • 34547400388 scopus 로고    scopus 로고
    • Evidence that long-term potentiation occurs within individual hippocampal synapses during learning
    • Fedulov V, Rex CS, Simmons DA, Palmer L, Gall CM, Lynch G (2007) Evidence that long-term potentiation occurs within individual hippocampal synapses during learning. J Neurosci 27:8031-8039.
    • (2007) J Neurosci , vol.27 , pp. 8031-8039
    • Fedulov, V.1    Rex, C.S.2    Simmons, D.A.3    Palmer, L.4    Gall, C.M.5    Lynch, G.6
  • 20
    • 0038662761 scopus 로고    scopus 로고
    • Hippocampal LTP is accompanied by enhanced F-actin content within the dendritic spine that is essential for late LTP maintenance in vivo
    • Fukazawa Y, Saitoh Y, Ozawa F, Ohta Y, Mizuno K, Inokuchi K (2003) Hippocampal LTP is accompanied by enhanced F-actin content within the dendritic spine that is essential for late LTP maintenance in vivo. Neuron 38:447-460.
    • (2003) Neuron , vol.38 , pp. 447-460
    • Fukazawa, Y.1    Saitoh, Y.2    Ozawa, F.3    Ohta, Y.4    Mizuno, K.5    Inokuchi, K.6
  • 21
    • 79955493637 scopus 로고    scopus 로고
    • Integrins traffic rapidly via circular dorsal ruffles and macropinocytosis during stimulated cell migration
    • Gu Z, Noss EH, Hsu VW, Brenner MB (2011) Integrins traffic rapidly via circular dorsal ruffles and macropinocytosis during stimulated cell migration. J Cell Biol 193:61-70.
    • (2011) J Cell Biol , vol.193 , pp. 61-70
    • Gu, Z.1    Noss, E.H.2    Hsu, V.W.3    Brenner, M.B.4
  • 22
    • 79251506941 scopus 로고    scopus 로고
    • HDAC inhibition modulates hippocampus-dependent longterm memory for object location in a CBP-dependent manner
    • Haettig J, Stefanko DP, Multani ML, Figueroa DX, McQuown SC, Wood MA (2011) HDAC inhibition modulates hippocampus-dependent longterm memory for object location in a CBP-dependent manner. Learn Mem 18:71-79.
    • (2011) Learn Mem , vol.18 , pp. 71-79
    • Haettig, J.1    Stefanko, D.P.2    Multani, M.L.3    Figueroa, D.X.4    McQuown, S.C.5    Wood, M.A.6
  • 23
    • 40049105259 scopus 로고    scopus 로고
    • The role of protein synthesis in memory consolidation: Progress amid decades of debate
    • Hernandez PJ, Abel T (2008) The role of protein synthesis in memory consolidation: progress amid decades of debate. Neurobiol Learn Mem 89:293-311.
    • (2008) Neurobiol Learn Mem , vol.89 , pp. 293-311
    • Hernandez, P.J.1    Abel, T.2
  • 24
    • 33751082371 scopus 로고    scopus 로고
    • Distinct roles of the beta 1-class integrins at the developing and the mature hippocampal excitatory synapse
    • Huang Z, Shimazu K, Woo NH, Zang K, Müller U, Lu B, Reichardt LF (2006) Distinct roles of the beta 1-class integrins at the developing and the mature hippocampal excitatory synapse. J Neurosci 26:11208-11219.
    • (2006) J Neurosci , vol.26 , pp. 11208-11219
    • Huang, Z.1    Shimazu, K.2    Woo, N.H.3    Zang, K.4    Müller, U.5    Lu, B.6    Reichardt, L.F.7
  • 26
    • 77957756704 scopus 로고    scopus 로고
    • Fibronectin and vitronectin induce AP-1-mediated matrix metalloproteinase-9 expression through integrin alpha(5)beta(1)/alpha(v)beta(3)-dependent Akt, ERK and JNK signaling pathways in human umbilical vein endothelial cells
    • Jin YJ, Park I, Hong IK, Byun HJ, Choi J, Kim YM, Lee H (2011) Fibronectin and vitronectin induce AP-1-mediated matrix metalloproteinase-9 expression through integrin alpha(5)beta(1)/alpha(v)beta(3)-dependent Akt, ERK and JNK signaling pathways in human umbilical vein endothelial cells. Cell Signal 23:125-134.
    • (2011) Cell Signal , vol.23 , pp. 125-134
    • Jin, Y.J.1    Park, I.2    Hong, I.K.3    Byun, H.J.4    Choi, J.5    Kim, Y.M.6    Lee, H.7
  • 27
    • 0026561901 scopus 로고
    • Brefeldin A: Insights into the control of membrane traffic and organelle structure
    • Klausner RD, Donaldson JG, Lippincott-Schwartz J (1992) Brefeldin A: insights into the control of membrane traffic and organelle structure. J Cell Biol 116:1071-1080.
    • (1992) J Cell Biol , vol.116 , pp. 1071-1080
    • Klausner, R.D.1    Donaldson, J.G.2    Lippincott-Schwartz, J.3
  • 28
    • 0036499748 scopus 로고    scopus 로고
    • Alpha3 integrin receptors contribute to the consolidation of long-term potentiation
    • Kramár EA, Bernard JA, Gall CM, Lynch G (2002) Alpha3 integrin receptors contribute to the consolidation of long-term potentiation. Neuroscience 110:29-39.
    • (2002) Neuroscience , vol.110 , pp. 29-39
    • Kramár, E.A.1    Bernard, J.A.2    Gall, C.M.3    Lynch, G.4
  • 29
  • 31
    • 0034612215 scopus 로고    scopus 로고
    • Dynamic actin filaments are required for stable long-term potentiation (LTP) in area CA1 of the hippocampus
    • Krucker T, Siggins GR, Halpain S (2000) Dynamic actin filaments are required for stable long-term potentiation (LTP) in area CA1 of the hippocampus. Proc Natl Acad Sci U S A 97:6856-6861.
    • (2000) Proc Natl Acad Sci U S A , vol.97 , pp. 6856-6861
    • Krucker, T.1    Siggins, G.R.2    Halpain, S.3
  • 32
    • 84855222932 scopus 로고    scopus 로고
    • The roles of the actin cytoskeleton in fear memory formation
    • Lamprecht R (2011) The roles of the actin cytoskeleton in fear memory formation. Front Behav Neurosci 5:39.
    • (2011) Front Behav Neurosci , vol.5 , pp. 39
    • Lamprecht, R.1
  • 33
    • 61449213806 scopus 로고    scopus 로고
    • Mechanisms that regulate adaptor binding to beta-integrin cytoplasmic tails
    • Legate KR, Fässler R (2009) Mechanisms that regulate adaptor binding to beta-integrin cytoplasmic tails. J Cell Sci 122:187-198.
    • (2009) J Cell Sci , vol.122 , pp. 187-198
    • Legate, K.R.1    Fässler, R.2
  • 35
    • 22244458242 scopus 로고    scopus 로고
    • AMPA receptor stimulation increases α5β1 integrin surface expression, adhesive function and signaling
    • Lin CY, Lynch G, Gall CM (2005) AMPA receptor stimulation increases α5β1 integrin surface expression, adhesive function and signaling. J Neurochem 94:531-546.
    • (2005) J Neurochem , vol.94 , pp. 531-546
    • Lin, C.Y.1    Lynch, G.2    Gall, C.M.3
  • 36
    • 41149130589 scopus 로고    scopus 로고
    • Integrin regulation of cytoplasmic calcium in excitatory neurons depends upon glutamate receptors and release from intracellular stores
    • Lin CY, Hilgenberg LG, Smith MA, Lynch G, Gall CM (2008) Integrin regulation of cytoplasmic calcium in excitatory neurons depends upon glutamate receptors and release from intracellular stores. Mol Cell Neurosci 37:770-780.
    • (2008) Mol Cell Neurosci , vol.37 , pp. 770-780
    • Lin, C.Y.1    Hilgenberg, L.G.2    Smith, M.A.3    Lynch, G.4    Gall, C.M.5
  • 37
    • 15844363687 scopus 로고    scopus 로고
    • Activated conformations of very late activation integrins detected by a group of antibodies (HUTS) specific for a novel regulatory region (355-425) of the common beta 1 chain
    • Luque A, Gómez M, Puzon W, Takada Y, Sánchez-Madrid F, Cabañas C (1996) Activated conformations of very late activation integrins detected by a group of antibodies (HUTS) specific for a novel regulatory region (355-425) of the common beta 1 chain. J Biol Chem 271:11067-11075.
    • (1996) J Biol Chem , vol.271 , pp. 11067-11075
    • Luque, A.1    Gómez, M.2    Puzon, W.3    Takada, Y.4    Sánchez-Madrid, F.5    Cabañas, C.6
  • 38
    • 33845863898 scopus 로고    scopus 로고
    • LTP consolidation: Substrates, explanatory power, and functional significance
    • Lynch G, Rex CS, Gall CM (2007) LTP consolidation: substrates, explanatory power, and functional significance. Neuropharmacology 52:12-23.
    • (2007) Neuropharmacology , vol.52 , pp. 12-23
    • Lynch, G.1    Rex, C.S.2    Gall, C.M.3
  • 39
    • 0346728801 scopus 로고    scopus 로고
    • Long-term potentiation and memory
    • Lynch MA (2004) Long-term potentiation and memory. Physiol Rev 84:87-136.
    • (2004) Physiol Rev , vol.84 , pp. 87-136
    • Lynch, M.A.1
  • 40
    • 0033969570 scopus 로고    scopus 로고
    • Memory-a century of consolidation
    • McGaugh JL (2000) Memory-a century of consolidation. Science 287:248-251.
    • (2000) Science , vol.287 , pp. 248-251
    • McGaugh, J.L.1
  • 41
    • 33748286819 scopus 로고    scopus 로고
    • Integrin-regulated FAK-Src signaling in normal and cancer cells
    • Mitra SK, Schlaepfer DD (2006) Integrin-regulated FAK-Src signaling in normal and cancer cells. Curr Opin Cell Biol 18:516-523.
    • (2006) Curr Opin Cell Biol , vol.18 , pp. 516-523
    • Mitra, S.K.1    Schlaepfer, D.D.2
  • 43
    • 84857200709 scopus 로고    scopus 로고
    • Differential involvement of protein synthesis and actin rearrangement in the reacquisition of contextual fear conditioning
    • Motanis H, Maroun M (2012) Differential involvement of protein synthesis and actin rearrangement in the reacquisition of contextual fear conditioning. Hippocampus 22:494-500.
    • (2012) Hippocampus , vol.22 , pp. 494-500
    • Motanis, H.1    Maroun, M.2
  • 44
    • 0002336051 scopus 로고
    • Experimentelle Beiträge zur Lehre vom Gedächtnis
    • Müller GE, Pilzecker A (1900) Experimentelle Beiträge zur Lehre vom Gedächtnis. Z Psychol S 1:1-288.
    • (1900) Z Psychol S , vol.1 , pp. 1-288
    • Müller, G.E.1    Pilzecker, A.2
  • 45
    • 33646552395 scopus 로고    scopus 로고
    • Reciprocal interactions between adhesion receptor signaling and MMP regulation
    • Munshi HG, Stack MS (2006) Reciprocal interactions between adhesion receptor signaling and MMP regulation. Cancer Metastasis Rev 25:45-56.
    • (2006) Cancer Metastasis Rev , vol.25 , pp. 45-56
    • Munshi, H.G.1    Stack, M.S.2
  • 47
    • 33646788001 scopus 로고    scopus 로고
    • Semaphorin 4D/Plexin-B1-mediated R-Ras GAP activity inhibits cell migration by regulating beta(1) integrin activity
    • Oinuma I, Katoh H, Negishi M (2006) Semaphorin 4D/Plexin-B1-mediated R-Ras GAP activity inhibits cell migration by regulating beta(1) integrin activity. J Cell Biol 173:601-613.
    • (2006) J Cell Biol , vol.173 , pp. 601-613
    • Oinuma, I.1    Katoh, H.2    Negishi, M.3
  • 48
    • 67650490390 scopus 로고    scopus 로고
    • Different Rho GTPase-dependent signaling pathways initiate sequential steps in the consolidation of long-term potentiation
    • Rex CS, Chen LY, Sharma A, Liu J, Babayan AH, Gall CM, Lynch G (2009) Different Rho GTPase-dependent signaling pathways initiate sequential steps in the consolidation of long-term potentiation. J Cell Biol 186:85-97.
    • (2009) J Cell Biol , vol.186 , pp. 85-97
    • Rex, C.S.1    Chen, L.Y.2    Sharma, A.3    Liu, J.4    Babayan, A.H.5    Gall, C.M.6    Lynch, G.7
  • 51
    • 41349089303 scopus 로고    scopus 로고
    • PKMzeta, LTP maintenance, and the dynamic molecular biology of memory storage
    • Sacktor TC (2008) PKMzeta, LTP maintenance, and the dynamic molecular biology of memory storage. Prog Brain Res 169:27-40.
    • (2008) Prog Brain Res , vol.169 , pp. 27-40
    • Sacktor, T.C.1
  • 52
    • 80053301137 scopus 로고    scopus 로고
    • Spatial and temporal regulation of integrin signalling during cell migration
    • Scales TM, Parsons M (2011) Spatial and temporal regulation of integrin signalling during cell migration. Curr Opin Cell Biol 23:562-568.
    • (2011) Curr Opin Cell Biol , vol.23 , pp. 562-568
    • Scales, T.M.1    Parsons, M.2
  • 53
    • 80051897106 scopus 로고    scopus 로고
    • Fibronectin-integrin (alpha5beta1) modulates migration and invasion of murine melanoma cell line B16F10 by involving MMP-9
    • Sil H, Sen T, Chatterjee A (2011) Fibronectin-integrin (alpha5beta1) modulates migration and invasion of murine melanoma cell line B16F10 by involving MMP-9. Oncol Res 19:335-348.
    • (2011) Oncol Res , vol.19 , pp. 335-348
    • Sil, H.1    Sen, T.2    Chatterjee, A.3
  • 54
    • 0034094370 scopus 로고    scopus 로고
    • Marking synaptic activity in dendritic spines with a calpain substrate exhibiting fluorescence resonance energy transfer
    • Vanderklish PW, Krushel LA, Holst BH, Gally JA, Crossin KL, Edelman GM (2000) Marking synaptic activity in dendritic spines with a calpain substrate exhibiting fluorescence resonance energy transfer. Proc Natl Acad Sci U S A 97:2253-2258.
    • (2000) Proc Natl Acad Sci U S A , vol.97 , pp. 2253-2258
    • Vanderklish, P.W.1    Krushel, L.A.2    Holst, B.H.3    Gally, J.A.4    Crossin, K.L.5    Edelman, G.M.6
  • 56
    • 58049191338 scopus 로고    scopus 로고
    • Extracellular proteolysis by matrix metalloproteinase-9 drives dendritic spine enlargement and long-term potentiation coordinately
    • Wang XB, Bozdagi O, Nikitczuk JS, Zhai ZW, Zhou Q, Huntley GW (2008) Extracellular proteolysis by matrix metalloproteinase-9 drives dendritic spine enlargement and long-term potentiation coordinately. Proc Natl Acad Sci U S A 105:19520-19525.
    • (2008) Proc Natl Acad Sci U S A , vol.105 , pp. 19520-19525
    • Wang, X.B.1    Bozdagi, O.2    Nikitczuk, J.S.3    Zhai, Z.W.4    Zhou, Q.5    Huntley, G.W.6
  • 57
    • 0009673621 scopus 로고
    • Single-trace fragility theory of memory dynamics
    • Wickelgren WA (1974) Single-trace fragility theory of memory dynamics. Mem Cognit 2:775-780.
    • (1974) Mem Cognit , vol.2 , pp. 775-780
    • Wickelgren, W.A.1
  • 58
    • 45949085855 scopus 로고    scopus 로고
    • Spine expansion and stabilization associated with long-term potentiation
    • Yang Y, Wang XB, Frerking M, Zhou Q (2008) Spine expansion and stabilization associated with long-term potentiation. J Neurosci 28:5740-5751.
    • (2008) J Neurosci , vol.28 , pp. 5740-5751
    • Yang, Y.1    Wang, X.B.2    Frerking, M.3    Zhou, Q.4


* 이 정보는 Elsevier사의 SCOPUS DB에서 KISTI가 분석하여 추출한 것입니다.