메뉴 건너뛰기




Volumn 7, Issue , 2016, Pages

Symmetric spike timing-dependent plasticity at CA3-CA3 synapses optimizes storage and recall in autoassociative networks

Author keywords

[No Author keywords available]

Indexed keywords

CELLS AND CELL COMPONENTS; CYTOGENETICS; GENE; MEMORY; NERVOUS SYSTEM; TEMPORAL PERIOD;

EID: 84968735621     PISSN: None     EISSN: 20411723     Source Type: Journal    
DOI: 10.1038/ncomms11552     Document Type: Article
Times cited : (130)

References (56)
  • 1
    • 36448992930 scopus 로고    scopus 로고
    • Behavioral functions of the CA3 subregion of the hippocampus
    • Kesner, R. P. Behavioral functions of the CA3 subregion of the hippocampus. Learn. Mem. 14, 771-781 (2007).
    • (2007) Learn. Mem. , vol.14 , pp. 771-781
    • Kesner, R.P.1
  • 3
    • 0020118274 scopus 로고
    • Neural networks and physical systems with emergent collective computational abilities
    • Hopfield, J. J. Neural networks and physical systems with emergent collective computational abilities. Proc. Natl Acad. Sci. USA 79, 2554-2558 (1982).
    • (1982) Proc. Natl Acad. Sci. USA , vol.79 , pp. 2554-2558
    • Hopfield, J.J.1
  • 4
    • 0023620017 scopus 로고
    • Hippocampal synaptic enhancement and information storage within a distributed memory system
    • McNaughton, B. L. & Morris, R. G. M. Hippocampal synaptic enhancement and information storage within a distributed memory system. Trends Neurosci. 10, 408-415 (1987).
    • (1987) Trends Neurosci. , vol.10 , pp. 408-415
    • McNaughton, B.L.1    Morris, R.G.M.2
  • 5
    • 0033082358 scopus 로고    scopus 로고
    • Relating hippocampal circuitry to function: Recall of memory sequences by reciprocal dentate-CA3 interactions
    • Lisman, J. E. Relating hippocampal circuitry to function: recall of memory sequences by reciprocal dentate-CA3 interactions. Neuron 22, 233-242 (1999).
    • (1999) Neuron , vol.22 , pp. 233-242
    • Lisman, J.E.1
  • 6
    • 0028109694 scopus 로고
    • Computational analysis of the role of the hippocampus in memory
    • Treves, A. & Rolls, E. T. Computational analysis of the role of the hippocampus in memory. Hippocampus 4, 374-391 (1994).
    • (1994) Hippocampus , vol.4 , pp. 374-391
    • Treves, A.1    Rolls, E.T.2
  • 7
    • 0028012436 scopus 로고
    • The hippocampal CA3 network: An in vivo intracellular labeling study
    • Li, X. G., Somogyi, P., Ylinen, A. & Buzsáki, G. The hippocampal CA3 network: an in vivo intracellular labeling study. J. Comp. Neurol. 339, 181-208 (1994).
    • (1994) J. Comp. Neurol. , vol.339 , pp. 181-208
    • Li, X.G.1    Somogyi, P.2    Ylinen, A.3    Buzsáki, G.4
  • 8
    • 0025352966 scopus 로고
    • Comparison of two forms of long-term potentiation in single hippocampal neurons
    • Zalutsky, R. A. & Nicoll, R. A. Comparison of two forms of long-term potentiation in single hippocampal neurons. Science 248, 1619-1624 (1990).
    • (1990) Science , vol.248 , pp. 1619-1624
    • Zalutsky, R.A.1    Nicoll, R.A.2
  • 9
    • 2642677679 scopus 로고    scopus 로고
    • Long-term synaptic plasticity between pairs of individual CA3 pyramidal cells in rat hippocampal slice cultures
    • Debanne, D., Gähwiler, B. H. & Thompson, S. M. Long-term synaptic plasticity between pairs of individual CA3 pyramidal cells in rat hippocampal slice cultures. J. Physiol. 507, 237-247 (1998).
    • (1998) J. Physiol. , vol.507 , pp. 237-247
    • Debanne, D.1    Gähwiler, B.H.2    Thompson, S.M.3
  • 10
    • 0035046162 scopus 로고    scopus 로고
    • Pair recordings reveal all-silent synaptic connections and the postsynaptic expression of long-term potentiation
    • Montgomery, J. M., Pavlidis, P. & Madison, D. V. Pair recordings reveal all-silent synaptic connections and the postsynaptic expression of long-term potentiation. Neuron 29, 691-701 (2001).
    • (2001) Neuron , vol.29 , pp. 691-701
    • Montgomery, J.M.1    Pavlidis, P.2    Madison, D.V.3
  • 11
    • 0037067583 scopus 로고    scopus 로고
    • Requirement for hippocampal CA3 NMDA receptors in associative memory recall
    • Nakazawa, K. et al. Requirement for hippocampal CA3 NMDA receptors in associative memory recall. Science 297, 211-218 (2002).
    • (2002) Science , vol.297 , pp. 211-218
    • Nakazawa, K.1
  • 12
    • 0026895844 scopus 로고
    • Statistical analysis of the dynamics of sparse associative memory
    • Gibson, W. G. & Robinson, J. Statistical analysis of the dynamics of sparse associative memory. Neural Networks 5, 645-661 (1992).
    • (1992) Neural Networks , vol.5 , pp. 645-661
    • Gibson, W.G.1    Robinson, J.2
  • 13
    • 0028781954 scopus 로고
    • Dynamics of the CA3 pyramidal neuron autoassociative memory network in the hippocampus
    • Bennett, M. R., Gibson, W. G. & Robinson, J. Dynamics of the CA3 pyramidal neuron autoassociative memory network in the hippocampus. Philos. Trans. R. Soc. Lond. B Biol. Sci. 343, 167-187 (1994).
    • (1994) Philos. Trans. R. Soc. Lond. B Biol. Sci. , vol.343 , pp. 167-187
    • Bennett, M.R.1    Gibson, W.G.2    Robinson, J.3
  • 15
    • 0031012390 scopus 로고    scopus 로고
    • A synaptically controlled, associative signal for Hebbian plasticity in hippocampal neurons
    • Magee, J. C. & Johnston, D. A synaptically controlled, associative signal for Hebbian plasticity in hippocampal neurons. Science 275, 209-213 (1997).
    • (1997) Science , vol.275 , pp. 209-213
    • Magee, J.C.1    Johnston, D.2
  • 16
    • 0031012615 scopus 로고    scopus 로고
    • Regulation of synaptic efficacy by coincidence of postsynaptic APs and EPSPs
    • Markram, H., Lübke, J., Frotscher, M. & Sakmann, B. Regulation of synaptic efficacy by coincidence of postsynaptic APs and EPSPs. Science 275, 213-215 (1997).
    • (1997) Science , vol.275 , pp. 213-215
    • Markram, H.1    Lübke, J.2    Frotscher, M.3    Sakmann, B.4
  • 17
    • 0032535029 scopus 로고    scopus 로고
    • Synaptic modifications in cultured hippocampal neurons: Dependence on spike timing, synaptic strength, and postsynaptic cell type
    • Bi, G.-Q. & Poo, M.-M. Synaptic modifications in cultured hippocampal neurons: Dependence on spike timing, synaptic strength, and postsynaptic cell type. J. Neurosci. 18, 10464-10472 (1998).
    • (1998) J. Neurosci. , vol.18 , pp. 10464-10472
    • Bi, G.-Q.1    Poo, M.-M.2
  • 18
    • 0034668762 scopus 로고    scopus 로고
    • 2+ channels
    • 2+ channels. J. Neurosci. 20, 8290-8297 (2000).
    • (2000) J. Neurosci. , vol.20 , pp. 8290-8297
    • Normann, C.1
  • 19
    • 0035924588 scopus 로고    scopus 로고
    • Rate, timing, and cooperativity jointly determine cortical synaptic plasticity
    • Sjöström, P. J., Turrigiano, G. G. & Nelson, S. B. Rate, timing, and cooperativity jointly determine cortical synaptic plasticity. Neuron 32, 1149-1164 (2001).
    • (2001) Neuron , vol.32 , pp. 1149-1164
    • Sjöström, P.J.1    Turrigiano, G.G.2    Nelson, S.B.3
  • 20
    • 84865401265 scopus 로고    scopus 로고
    • The spike-timing dependence of plasticity
    • Feldman, D. E. The spike-timing dependence of plasticity. Neuron 75, 556-571 (2012).
    • (2012) Neuron , vol.75 , pp. 556-571
    • Feldman, D.E.1
  • 21
    • 84862793600 scopus 로고    scopus 로고
    • Active dendrites support efficient initiation of dendritic spikes in hippocampal CA3 pyramidal neurons
    • Kim, S., Guzman, S. J., Hu, H. & Jonas, P. Active dendrites support efficient initiation of dendritic spikes in hippocampal CA3 pyramidal neurons. Nat. Neurosci. 15, 600-606 (2012).
    • (2012) Nat. Neurosci , vol.15 , pp. 600-606
    • Kim, S.1    Guzman, S.J.2    Hu, H.3    Jonas, P.4
  • 22
    • 79952123974 scopus 로고    scopus 로고
    • Axonal morphometry of hippocampal pyramidal neurons semi-automatically reconstructed after in vivo labeling in different CA3 locations
    • Ropireddy, D., Scorcioni, R., Lasher, B., Buzsáki, G. & Ascoli, G. A. Axonal morphometry of hippocampal pyramidal neurons semi-automatically reconstructed after in vivo labeling in different CA3 locations. Brain Struct. Funct. 216, 1-15 (2011).
    • (2011) Brain Struct. Funct. , vol.216 , pp. 1-15
    • Ropireddy, D.1    Scorcioni, R.2    Lasher, B.3    Buzsáki, G.4    Ascoli, G.A.5
  • 23
    • 85057293939 scopus 로고    scopus 로고
    • Properties of CA3-CA3 synapses optimize storage capacity in the CA3 microcircuit
    • Guzman, S. J., Frotscher, M. & Jonas, P. Properties of CA3-CA3 synapses optimize storage capacity in the CA3 microcircuit. Soc. of Neurosci. Abstr. 302, 11/C33 (2014).
    • (2014) Soc. of Neurosci. Abstr. , vol.302 , pp. 11/C33
    • Guzman, S.J.1    Frotscher, M.2    Jonas, P.3
  • 24
    • 15044340596 scopus 로고    scopus 로고
    • Spike-timing-dependent synaptic plasticity depends on dendritic location
    • Froemke, R. C., Poo, M. M. & Dan, Y. Spike-timing-dependent synaptic plasticity depends on dendritic location. Nature 434, 221-225 (2005).
    • (2005) Nature , vol.434 , pp. 221-225
    • Froemke, R.C.1    Poo, M.M.2    Dan, Y.3
  • 25
    • 33749855263 scopus 로고    scopus 로고
    • Learning rules for spike timing-dependent plasticity depend on dendritic synapse location
    • Letzkus, J. J., Kampa, B. M. & Stuart, G. J. Learning rules for spike timing-dependent plasticity depend on dendritic synapse location. J. Neurosci. 26, 10420-10429 (2006).
    • (2006) J. Neurosci. , vol.26 , pp. 10420-10429
    • Letzkus, J.J.1    Kampa, B.M.2    Stuart, G.J.3
  • 26
    • 33745902544 scopus 로고    scopus 로고
    • A cooperative switch determines the sign of synaptic plasticity in distal dendrites of neocortical pyramidal neurons
    • Sjöström, P. J. & Häusser, M. A cooperative switch determines the sign of synaptic plasticity in distal dendrites of neocortical pyramidal neurons. Neuron 51, 227-238 (2006).
    • (2006) Neuron , vol.51 , pp. 227-238
    • Sjöström, P.J.1    Häusser, M.2
  • 27
    • 62749207175 scopus 로고    scopus 로고
    • Activity-dependent depression of the spike after-depolarization generates long-lasting intrinsic plasticity in hippocampal CA3 pyramidal neurons
    • Brown, J. T. & Randall, A. D. Activity-dependent depression of the spike after-depolarization generates long-lasting intrinsic plasticity in hippocampal CA3 pyramidal neurons. J. Physiol. 587, 1265-1281 (2009).
    • (2009) J. Physiol. , vol.587 , pp. 1265-1281
    • Brown, J.T.1    Randall, A.D.2
  • 28
    • 20544443789 scopus 로고    scopus 로고
    • R-type calcium channels contribute to afterdepolarization and bursting in hippocampal CA1 pyramidal neurons
    • Metz, A. E., Jarsky, T., Martina, M. & Spruston, N. R-type calcium channels contribute to afterdepolarization and bursting in hippocampal CA1 pyramidal neurons. J. Neurosci. 25, 5763-5773 (2005).
    • (2005) J. Neurosci. , vol.25 , pp. 5763-5773
    • Metz, A.E.1    Jarsky, T.2    Martina, M.3    Spruston, N.4
  • 29
    • 34248224617 scopus 로고    scopus 로고
    • Dendritic D-type potassium currents inhibit the spike afterdepolarization in rat hippocampal CA1 pyramidal neurons
    • Metz, A. E., Spruston, N. & Martina, M. Dendritic D-type potassium currents inhibit the spike afterdepolarization in rat hippocampal CA1 pyramidal neurons. J. Physiol. 15, 175-187 (2007).
    • (2007) J. Physiol. , vol.15 , pp. 175-187
    • Metz, A.E.1    Spruston, N.2    Martina, M.3
  • 30
    • 0029899046 scopus 로고    scopus 로고
    • Functional significance of long-term potentiation for sequence learning and prediction
    • Abbott, L. F. & Blum, K. I. Functional significance of long-term potentiation for sequence learning and prediction. Cereb. Cortex 6, 406-416 (1996).
    • (1996) Cereb. Cortex , vol.6 , pp. 406-416
    • Abbott, L.F.1    Blum, K.I.2
  • 31
    • 0033725001 scopus 로고    scopus 로고
    • Reversible associative depression and nonassociative potentiation at a parallel fiber synapse
    • Han, V. Z., Grant, K. & Bell, C. C. Reversible associative depression and nonassociative potentiation at a parallel fiber synapse. Neuron 27, 611-622 (2000).
    • (2000) Neuron , vol.27 , pp. 611-622
    • Han, V.Z.1    Grant, K.2    Bell, C.C.3
  • 32
    • 2542552033 scopus 로고    scopus 로고
    • Firing mode-dependent synaptic plasticity in rat neocortical pyramidal neurons
    • Birtoli, B. & Ulrich, D. Firing mode-dependent synaptic plasticity in rat neocortical pyramidal neurons. J. Neurosci. 24, 4935-4940 (2004).
    • (2004) J. Neurosci. , vol.24 , pp. 4935-4940
    • Birtoli, B.1    Ulrich, D.2
  • 33
    • 30544442754 scopus 로고    scopus 로고
    • Bidirectional activity-dependent plasticity at corticostriatal synapses
    • Fino, E., Glowinski, J. & Venance, L. Bidirectional activity-dependent plasticity at corticostriatal synapses. J. Neurosci. 25, 11279-11287 (2005).
    • (2005) J. Neurosci. , vol.25 , pp. 11279-11287
    • Fino, E.1    Glowinski, J.2    Venance, L.3
  • 34
    • 34848871579 scopus 로고    scopus 로고
    • Spike-timing-dependent plasticity of neocortical excitatory synapses on inhibitory interneurons depends on target cell type
    • Lu, J. T., Li, C. Y., Zhao, J. P., Poo, M. M. & Zhang, X. H. Spike-timing-dependent plasticity of neocortical excitatory synapses on inhibitory interneurons depends on target cell type. J. Neurosci. 27, 9711-9720 (2007).
    • (2007) J. Neurosci. , vol.27 , pp. 9711-9720
    • Lu, J.T.1    Li, C.Y.2    Zhao, J.P.3    Poo, M.M.4    Zhang, X.H.5
  • 35
    • 37349086109 scopus 로고    scopus 로고
    • Timing dependence of the induction of cerebellar LTD
    • Safo, P. & Regehr, W. G. Timing dependence of the induction of cerebellar LTD. Neuropharmacology 54, 213-218 (2008).
    • (2008) Neuropharmacology , vol.54 , pp. 213-218
    • Safo, P.1    Regehr, W.G.2
  • 36
    • 84906057141 scopus 로고    scopus 로고
    • Depression biased non-Hebbian spike-timing-dependent synaptic plasticity in the rat subiculum
    • Pandey, A. & Sikdar, S. K. Depression biased non-Hebbian spike-timing-dependent synaptic plasticity in the rat subiculum. J. Physiol. 592, 3537-3557 (2014).
    • (2014) J. Physiol. , vol.592 , pp. 3537-3557
    • Pandey, A.1    Sikdar, S.K.2
  • 37
    • 33751119889 scopus 로고    scopus 로고
    • 2+ signaling in spike-timing-dependent plasticity
    • 2+ signaling in spike-timing-dependent plasticity. J. Neurosci. 26, 11001-11013 (2006).
    • (2006) J. Neurosci. , vol.26 , pp. 11001-11013
    • Nevian, T.1    Sakmann, B.2
  • 38
    • 34548503919 scopus 로고    scopus 로고
    • Dendritic mechanisms controlling spike-timing-dependent synaptic plasticity
    • Kampa, B. M., Letzkus, J. J. & Stuart, G. J. Dendritic mechanisms controlling spike-timing-dependent synaptic plasticity. Trends Neurosci. 30, 456-463 (2007).
    • (2007) Trends Neurosci. , vol.30 , pp. 456-463
    • Kampa, B.M.1    Letzkus, J.J.2    Stuart, G.J.3
  • 40
    • 2342467434 scopus 로고    scopus 로고
    • 2+ unblock of NMDA receptors: Implications for spike-timing dependent synaptic plasticity
    • 2+ unblock of NMDA receptors: implications for spike-timing dependent synaptic plasticity. J. Physiol. 556, 337-345 (2004).
    • (2004) J. Physiol. , vol.556 , pp. 337-345
    • Kampa, B.M.1    Clements, J.2    Jonas, P.3    Stuart, G.J.4
  • 41
    • 0035808602 scopus 로고    scopus 로고
    • Differential shunting of EPSPs by action potentials
    • Häusser, M., Major, G. & Stuart, G. J. Differential shunting of EPSPs by action potentials. Science 291, 138-141 (2011).
    • (2011) Science , vol.291 , pp. 138-141
    • Häusser, M.1    Major, G.2    Stuart, G.J.3
  • 42
    • 0035155356 scopus 로고    scopus 로고
    • Dendritic coincidence detection of EPSPs and action potentials
    • Stuart, G. J. & Häusser, M. Dendritic coincidence detection of EPSPs and action potentials. Nat. Neurosci. 4, 63-71 (2001).
    • (2001) Nat. Neurosci , vol.4 , pp. 63-71
    • Stuart, G.J.1    Häusser, M.2
  • 43
    • 84907336769 scopus 로고    scopus 로고
    • Synapse-specific control of experience-dependent plasticity by presynaptic NMDA receptors
    • Larsen, R. S. et al. Synapse-specific control of experience-dependent plasticity by presynaptic NMDA receptors. Neuron 83, 879-893 (2014).
    • (2014) Neuron , vol.83 , pp. 879-893
    • Larsen, R.S.1
  • 44
    • 84946600641 scopus 로고    scopus 로고
    • Retroactive modulation of spike timing-dependent plasticity by dopamine
    • Brzosko, Z., Schultz, W. & Paulsen, O. Retroactive modulation of spike timing-dependent plasticity by dopamine. Elife 4, e09685 (2015).
    • (2015) Elife , vol.4
    • Brzosko, Z.1    Schultz, W.2    Paulsen, O.3
  • 45
    • 0742323527 scopus 로고    scopus 로고
    • Homeostatic plasticity in the developing nervous system
    • Turrigiano, G. G. & Nelson, S. B. Homeostatic plasticity in the developing nervous system. Nat. Rev. Neurosci. 5, 97-107 (2004).
    • (2004) Nat. Rev. Neurosci. , vol.5 , pp. 97-107
    • Turrigiano, G.G.1    Nelson, S.B.2
  • 46
    • 84856259585 scopus 로고    scopus 로고
    • Heterogeneous reallocation of presynaptic efficacy in recurrent excitatory circuits adapting to inactivity
    • Mitra, A., Mitra, S. S. & Tsien, R. W. Heterogeneous reallocation of presynaptic efficacy in recurrent excitatory circuits adapting to inactivity. Nat. Neurosci. 15, 250-257 (2011).
    • (2011) Nat. Neurosci , vol.15 , pp. 250-257
    • Mitra, A.1    Mitra, S.S.2    Tsien, R.W.3
  • 47
    • 84872175082 scopus 로고    scopus 로고
    • Mossy fiber-CA3 synapses mediate homeostatic plasticity in mature hippocampal neurons
    • Lee, K. J. et al. Mossy fiber-CA3 synapses mediate homeostatic plasticity in mature hippocampal neurons. Neuron 77, 99-114 (2013).
    • (2013) Neuron , vol.77 , pp. 99-114
    • Lee, K.J.1
  • 48
    • 0242524496 scopus 로고    scopus 로고
    • Depolarization-induced long-term depression at hippocampal mossy fiber-CA3 pyramidal neuron synapses
    • Lei, S. et al. Depolarization-induced long-term depression at hippocampal mossy fiber-CA3 pyramidal neuron synapses. J. Neurosci. 23, 9786-9795 (2003).
    • (2003) J. Neurosci. , vol.23 , pp. 9786-9795
    • Lei, S.1
  • 49
    • 35548940818 scopus 로고    scopus 로고
    • 2+-permeable AMPA receptors underlies depolarization-induced long-term depression at mossy fiber CA3 pyramid synapses
    • 2+-permeable AMPA receptors underlies depolarization-induced long-term depression at mossy fiber CA3 pyramid synapses. J. Neurosci. 27, 11651-11662 (2007).
    • (2007) J. Neurosci. , vol.27 , pp. 11651-11662
    • Ho, M.T.1
  • 50
    • 38649095256 scopus 로고    scopus 로고
    • Reactivation of experience-dependent cell assembly patterns in the hippocampus
    • O'Neill, J., Senior, T. J., Allen, K., Huxter, J. R. & Csicsvari, J. Reactivation of experience-dependent cell assembly patterns in the hippocampus. Nat. Neurosci. 11, 209-215 (2008).
    • (2008) Nat. Neurosci , vol.11 , pp. 209-215
    • O'Neill, J.1    Senior, T.J.2    Allen, K.3    Huxter, J.R.4    Csicsvari, J.5
  • 51
    • 0022557253 scopus 로고
    • Excitatory synaptic interactions between CA3 neurones in the guinea-pig hippocampus
    • Miles, R. & Wong, R. K. S. Excitatory synaptic interactions between CA3 neurones in the guinea-pig hippocampus. J. Physiol. 373, 397-418 (1986).
    • (1986) J. Physiol. , vol.373 , pp. 397-418
    • Miles, R.1    Wong, R.K.S.2
  • 52
    • 0029995540 scopus 로고    scopus 로고
    • Activation of metabotropic glutamate receptor type 2/3 suppresses transmission at rat hippocampal mossy fibre synapses
    • Kamiya, H., Shinozaki, H. & Yamamoto, C. Activation of metabotropic glutamate receptor type 2/3 suppresses transmission at rat hippocampal mossy fibre synapses. J. Physiol. 493, 447-455 (1996).
    • (1996) J. Physiol. , vol.493 , pp. 447-455
    • Kamiya, H.1    Shinozaki, H.2    Yamamoto, C.3
  • 53
    • 0036303258 scopus 로고    scopus 로고
    • Facilitation at single synapses probed with optical quantal analysis
    • Oertner, T. G., Sabatini, B. L., Nimchinsky, E. & Svoboda, K. Facilitation at single synapses probed with optical quantal analysis. Nat. Neurosci. 5, 657-664 (2002).
    • (2002) Nat. Neurosci , vol.5 , pp. 657-664
    • Oertner, T.G.1    Sabatini, B.L.2    Nimchinsky, E.3    Svoboda, K.4
  • 54
    • 84862520770 scopus 로고    scopus 로고
    • Fiji: An open-source platform for biological-image analysis
    • Schindelin, J. et al. Fiji: an open-source platform for biological-image analysis. Nat. Methods 9, 676-682 (2012).
    • (2012) Nat. Methods , vol.9 , pp. 676-682
    • Schindelin, J.1
  • 56
    • 33745712258 scopus 로고    scopus 로고
    • A review of the integrate-and-fire neuron model: I. Homogeneous synaptic input
    • Burkitt, A. N. A review of the integrate-and-fire neuron model: I. Homogeneous synaptic input. Biol. Cybern. 95, 1-19 (2006).
    • (2006) Biol. Cybern. , vol.95 , pp. 1-19
    • Burkitt, A.N.1


* 이 정보는 Elsevier사의 SCOPUS DB에서 KISTI가 분석하여 추출한 것입니다.