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Volumn 11, Issue 3, 2014, Pages 338-346

Independent optical excitation of distinct neural populations

(21)  Klapoetke, Nathan C a,b,c,d   Murata, Yasunobu c,d   Kim, Sung Soo e   Pulver, Stefan R e   Birdsey Benson, Amanda c,d   Cho, Yong Ku a,b,c,d   Morimoto, Tania K a,b,c,d   Chuong, Amy S a,b,c,d   Carpenter, Eric J f   Tian, Zhijian g   Wang, Jun g   Xie, Yinlong g   Yan, Zhixiang g   Zhang, Yong g   Chow, Brian Y h   Surek, Barbara i   Melkonian, Michael i   Jayaraman, Vivek e   Constantine Paton, Martha c,d   Wong, Gane Ka Shu f,g   more..


Author keywords

[No Author keywords available]

Indexed keywords

CHRIMSON; CHRONOS; OPSIN; RHODOPSIN; UNCLASSIFIED DRUG;

EID: 84895524488     PISSN: 15487091     EISSN: 15487105     Source Type: Journal    
DOI: 10.1038/nmeth.2836     Document Type: Article
Times cited : (1569)

References (50)
  • 1
    • 26444621497 scopus 로고    scopus 로고
    • Millisecond-timescale genetically targeted optical control of neural activity
    • Boyden, E.S., Zhang, F., Bamberg, E., Nagel, G. & Deisseroth, K. Millisecond-timescale, genetically targeted optical control of neural activity. Nat. Neurosci. 8, 1263-1268 (2005).
    • (2005) Nat. Neurosci. , vol.8 , pp. 1263-1268
    • Boyden, E.S.1    Zhang, F.2    Bamberg, E.3    Nagel, G.4    Deisseroth, K.5
  • 2
    • 55849136898 scopus 로고    scopus 로고
    • Multiple-color optical activation, silencing, and desynchronization of neural activity, with single-spike temporal resolution
    • Han, X. & Boyden, E.S. Multiple-color optical activation, silencing, and desynchronization of neural activity, with single-spike temporal resolution. PLoS ONE 2, e299 (2007).
    • (2007) PLoS ONE , vol.2
    • Han, X.1    Boyden, E.S.2
  • 3
    • 73849143150 scopus 로고    scopus 로고
    • High-performance genetically targetable optical neural silencing by light-driven proton pumps
    • Chow, B.Y. et al. High-performance genetically targetable optical neural silencing by light-driven proton pumps. Nature 463, 98-102 (2010).
    • (2010) Nature , vol.463 , pp. 98-102
    • Chow, B.Y.1
  • 4
    • 34247222730 scopus 로고    scopus 로고
    • Multimodal fast optical interrogation of neural circuitry
    • Zhang, F. et al. Multimodal fast optical interrogation of neural circuitry. Nature 446, 633-639 (2007).
    • (2007) Nature , vol.446 , pp. 633-639
    • Zhang, F.1
  • 5
    • 77950930288 scopus 로고    scopus 로고
    • Molecular and cellular approaches for diversifying and extending optogenetics
    • Gradinaru, V. et al. Molecular and cellular approaches for diversifying and extending optogenetics. Cell 141, 154-165 (2010).
    • (2010) Cell , vol.141 , pp. 154-165
    • Gradinaru, V.1
  • 6
    • 79955684899 scopus 로고    scopus 로고
    • A history of optogenetics: The development of tools for controlling brain circuits with light
    • Boyden, E.S. A history of optogenetics: the development of tools for controlling brain circuits with light. F1000 Biol. Rep. 3, 11 (2011).
    • (2011) F1000 Biol. Rep. , vol.3 , pp. 11
    • Boyden, E.S.1
  • 7
    • 44349120639 scopus 로고    scopus 로고
    • Red-shifted optogenetic excitation: A tool for fast neural control derived from Volvox carteri
    • Zhang, F. et al. Red-shifted optogenetic excitation: a tool for fast neural control derived from Volvox carteri. Nat. Neurosci. 11, 631-633 (2008).
    • (2008) Nat. Neurosci. , vol.11 , pp. 631-633
    • Zhang, F.1
  • 8
    • 84867077532 scopus 로고    scopus 로고
    • Bimodal activation of different neuron classes with the spectrally red-shifted channelrhodopsin chimera C1V1 in Caenorhabditis elegans
    • Erbguth, K., Prigge, M., Schneider, F., Hegemann, P. & Gottschalk, A. Bimodal activation of different neuron classes with the spectrally red-shifted channelrhodopsin chimera C1V1 in Caenorhabditis elegans. PLoS ONE 7, e46827 (2012).
    • (2012) PLoS ONE , vol.7
    • Erbguth, K.1    Prigge, M.2    Schneider, F.3    Hegemann, P.4    Gottschalk, A.5
  • 9
    • 80052563319 scopus 로고    scopus 로고
    • Neocortical excitation/inhibition balance in information processing and social dysfunction
    • Yizhar, O. et al. Neocortical excitation/inhibition balance in information processing and social dysfunction. Nature 477, 171-178 (2011).
    • (2011) Nature , vol.477 , pp. 171-178
    • Yizhar, O.1
  • 10
    • 84866434885 scopus 로고    scopus 로고
    • Color-tuned channelrhodopsins for multiwavelength optogenetics
    • Prigge, M. et al. Color-tuned channelrhodopsins for multiwavelength optogenetics. J. Biol. Chem. 287, 31804-31812 (2012).
    • (2012) J. Biol. Chem. , vol.287 , pp. 31804-31812
    • Prigge, M.1
  • 11
    • 84870696480 scopus 로고    scopus 로고
    • Tuning the electronic absorption of protein-embedded all-trans-retinal
    • Wang, W. et al. Tuning the electronic absorption of protein-embedded all-trans-retinal. Science 338, 1340-1343 (2012).
    • (2012) Science , vol.338 , pp. 1340-1343
    • Wang, W.1
  • 12
    • 0015921932 scopus 로고
    • Visual pigments determination and interpretation of the fluorescence quantum yields of retinals, Schiff bases, and protonated Schiff bases
    • Waddell, W.H., Schaffer, A.M. & Becker, R.S. Visual pigments Determination and interpretation of the fluorescence quantum yields of retinals, Schiff bases, and protonated Schiff bases. J. Am. Chem. Soc. 95, 8223-8227 (1973).
    • (1973) J. Am. Chem. Soc. , vol.95 , pp. 8223-8227
    • Waddell, W.H.1    Schaffer, A.M.2    Becker, R.S.3
  • 13
    • 84855190225 scopus 로고    scopus 로고
    • New channelrhodopsin with a red-shifted spectrum and rapid kinetics from Mesostigma viride
    • Govorunova, E.G., Spudich, E.N., Lane, C.E., Sineshchekov, O.A. & Spudich, J.L. New channelrhodopsin with a red-shifted spectrum and rapid kinetics from Mesostigma viride. mBio 2, e00115-e00111 (2011).
    • (2011) MBio , vol.2
    • Govorunova, E.G.1    Spudich, E.N.2    Lane, C.E.3    Sineshchekov, O.A.4    Spudich, J.L.5
  • 14
    • 84884906500 scopus 로고    scopus 로고
    • ReaChR: A red-shifted variant of channelrhodopsin enables deep transcranial optogenetic excitation
    • Lin, J.Y., Knutsen, P.M., Muller, A., Kleinfeld, D. & Tsien, R.Y. ReaChR: a red-shifted variant of channelrhodopsin enables deep transcranial optogenetic excitation. Nat. Neurosci. 16, 1499-1508 (2013).
    • (2013) Nat. Neurosci. , vol.16 , pp. 1499-1508
    • Lin, J.Y.1    Knutsen, P.M.2    Muller, A.3    Kleinfeld, D.4    Tsien, R.Y.5
  • 15
    • 79953230429 scopus 로고    scopus 로고
    • Ultra light-sensitive and fast neuronal activation with the Ca2+-permeable channelrhodopsin CatCh
    • Kleinlogel, S. et al. Ultra light-sensitive and fast neuronal activation with the Ca2+-permeable channelrhodopsin CatCh. Nat. Neurosci. 14, 513-518 (2011).
    • (2011) Nat. Neurosci. , vol.14 , pp. 513-518
    • Kleinlogel, S.1
  • 17
    • 75349085542 scopus 로고    scopus 로고
    • Structural guidance of the photocycle of channelrhodopsin-2 by an interhelical hydrogen bond
    • Bamann, C., Gueta, R., Kleinlogel, S., Nagel, G. & Bamberg, E. Structural guidance of the photocycle of channelrhodopsin-2 by an interhelical hydrogen bond. Biochemistry 49, 267-278 (2010).
    • (2010) Biochemistry , vol.49 , pp. 267-278
    • Bamann, C.1    Gueta, R.2    Kleinlogel, S.3    Nagel, G.4    Bamberg, E.5
  • 18
    • 84856502824 scopus 로고    scopus 로고
    • Principles for applying optogenetic tools derived from direct comparative analysis of microbial opsins
    • Mattis, J. et al. Principles for applying optogenetic tools derived from direct comparative analysis of microbial opsins. Nat. Methods 9, 159-172 (2012).
    • (2012) Nat. Methods , vol.9 , pp. 159-172
    • Mattis, J.1
  • 19
    • 84885605921 scopus 로고    scopus 로고
    • Characterization of a highly efficient blue-shifted channelrhodopsin from the marine alga Platymonas subcordiformis
    • Govorunova, E.G., Sineshchekov, O.A., Li, H., Janz, R. & Spudich, J.L. Characterization of a highly efficient blue-shifted channelrhodopsin from the marine alga Platymonas subcordiformis. J. Biol. Chem. 288, 29911-29922 (2013).
    • (2013) J. Biol. Chem. , vol.288 , pp. 29911-29922
    • Govorunova, E.G.1    Sineshchekov, O.A.2    Li, H.3    Janz, R.4    Spudich, J.L.5
  • 20
    • 84869751747 scopus 로고    scopus 로고
    • Evaluating methods for isolating total RNA and predicting the success of sequencing phylogenetically diverse plant transcriptomes
    • Johnson, M.T. et al. Evaluating methods for isolating total RNA and predicting the success of sequencing phylogenetically diverse plant transcriptomes. PLoS ONE 7, e50226 (2012).
    • (2012) PLoS ONE , vol.7
    • Johnson, M.T.1
  • 21
    • 78049261560 scopus 로고    scopus 로고
    • A user's guide to channelrhodopsin variants: Features, limitations and future developments
    • Lin, J.Y. A user's guide to channelrhodopsin variants: features, limitations and future developments. Exp. Physiol. 96, 19-25 (2011).
    • (2011) Exp. Physiol. , vol.96 , pp. 19-25
    • Lin, J.Y.1
  • 22
    • 29044433616 scopus 로고    scopus 로고
    • Light activation of channelrhodopsin-2 in excitable cells of Caenorhabditis elegans triggers rapid behavioral responses
    • Nagel, G. et al. Light activation of channelrhodopsin-2 in excitable cells of Caenorhabditis elegans triggers rapid behavioral responses. Curr. Biol. 15, 2279-2284 (2005).
    • (2005) Curr. Biol. , vol.15 , pp. 2279-2284
    • Nagel, G.1
  • 23
    • 84857933789 scopus 로고    scopus 로고
    • Loom-sensitive neurons link computation to action in the Drosophila visual system
    • de Vries, S.E. & Clandinin, T.R. Loom-sensitive neurons link computation to action in the Drosophila visual system. Curr. Biol. 22, 353-362 (2012).
    • (2012) Curr. Biol. , vol.22 , pp. 353-362
    • De Vries, S.E.1    Clandinin, T.R.2
  • 24
    • 33749837676 scopus 로고    scopus 로고
    • Light-induced activation of distinct modulatory neurons triggers appetitive or aversive learning in Drosophila larvae
    • Schroll, C. et al. Light-induced activation of distinct modulatory neurons triggers appetitive or aversive learning in Drosophila larvae. Curr. Biol. 16, 1741-1747 (2006).
    • (2006) Curr. Biol. , vol.16 , pp. 1741-1747
    • Schroll, C.1
  • 25
    • 84872607103 scopus 로고    scopus 로고
    • Asymmetric neurotransmitter release enables rapid odour lateralization in Drosophila
    • Gaudry, Q., Hong, E.J., Kain, J., de Bivort, B.L. & Wilson, R.I. Asymmetric neurotransmitter release enables rapid odour lateralization in Drosophila. Nature 493, 424-428 (2013).
    • (2013) Nature , vol.493 , pp. 424-428
    • Gaudry, Q.1    Hong, E.J.2    Kain, J.3    De Bivort, B.L.4    Wilson, R.I.5
  • 26
    • 84864430036 scopus 로고    scopus 로고
    • Optogenetic manipulation of neural circuits and behavior in Drosophila larvae
    • Honjo, K., Hwang, R.Y. & Tracey, W.D. Jr. Optogenetic manipulation of neural circuits and behavior in Drosophila larvae. Nat. Protoc. 7, 1470-1478 (2012).
    • (2012) Nat. Protoc. , vol.7 , pp. 1470-1478
    • Honjo, K.1    Hwang, R.Y.2    Tracey Jr., W.D.3
  • 27
    • 35748963218 scopus 로고    scopus 로고
    • A toolbox for light control of Drosophila behaviors through Channelrhodopsin 2-mediated photoactivation of targeted neurons
    • Zhang, W., Ge, W. & Wang, Z. A toolbox for light control of Drosophila behaviors through Channelrhodopsin 2-mediated photoactivation of targeted neurons. Eur. J. Neurosci. 26, 2405-2416 (2007).
    • (2007) Eur. J. Neurosci. , vol.26 , pp. 2405-2416
    • Zhang, W.1    Ge, W.2    Wang, Z.3
  • 28
    • 78650306355 scopus 로고    scopus 로고
    • Light-avoidance-mediating photoreceptors tile the Drosophila larval body wall
    • Xiang, Y. et al. Light-avoidance-mediating photoreceptors tile the Drosophila larval body wall. Nature 468, 921-926 (2010).
    • (2010) Nature , vol.468 , pp. 921-926
    • Xiang, Y.1
  • 29
    • 84895558645 scopus 로고    scopus 로고
    • Optogenetic control of Drosophila using a red-shifted channelrhodopsin reveals experience-dependent influences on courtship
    • doi:10.1038/nmeth.2765 22 December 2013
    • Inagaki, H.K. et al. Optogenetic control of Drosophila using a red-shifted channelrhodopsin reveals experience-dependent influences on courtship. Nat. Methods doi:10.1038/nmeth.2765 (22 December 2013).
    • Nat. Methods
    • Inagaki, H.K.1
  • 30
    • 70349804300 scopus 로고    scopus 로고
    • Writing memories with light-addressable reinforcement circuitry
    • Claridge-Chang, A. et al. Writing memories with light-addressable reinforcement circuitry. Cell 139, 405-415 (2009).
    • (2009) Cell , vol.139 , pp. 405-415
    • Claridge-Chang, A.1
  • 31
    • 66349129337 scopus 로고    scopus 로고
    • Temporal dynamics of neuronal activation by Channelrhodopsin-2 and TRPA1 determine behavioral output in Drosophila larvae
    • Pulver, S.R., Pashkovski, S.L., Hornstein, N.J., Garrity, P.A. & Griffith, L.C. Temporal dynamics of neuronal activation by Channelrhodopsin-2 and TRPA1 determine behavioral output in Drosophila larvae. J. Neurophysiol. 101, 3075-3088 (2009).
    • (2009) J. Neurophysiol. , vol.101 , pp. 3075-3088
    • Pulver, S.R.1    Pashkovski, S.L.2    Hornstein, N.J.3    Garrity, P.A.4    Griffith, L.C.5
  • 32
    • 84857686193 scopus 로고    scopus 로고
    • Optogenetics and thermogenetics: Technologies for controlling the activity of targeted cells within intact neural circuits
    • Bernstein, J.G., Garrity, P.A. & Boyden, E.S. Optogenetics and thermogenetics: technologies for controlling the activity of targeted cells within intact neural circuits. Curr. Opin. Neurobiol. 22, 61-71 (2012).
    • (2012) Curr. Opin. Neurobiol. , vol.22 , pp. 61-71
    • Bernstein, J.G.1    Garrity, P.A.2    Boyden, E.S.3
  • 33
    • 36849017592 scopus 로고    scopus 로고
    • Nociceptive neurons protect Drosophila larvae from parasitoid wasps
    • Hwang, R.Y. et al. Nociceptive neurons protect Drosophila larvae from parasitoid wasps. Curr. Biol. 17, 2105-2116 (2007).
    • (2007) Curr. Biol. , vol.17 , pp. 2105-2116
    • Hwang, R.Y.1
  • 34
    • 35648936454 scopus 로고    scopus 로고
    • Two Gr genes underlie sugar reception in Drosophila
    • Dahanukar, A., Lei, Y.T., Kwon, J.Y. & Carlson, J.R. Two Gr genes underlie sugar reception in Drosophila. Neuron 56, 503-516 (2007).
    • (2007) Neuron , vol.56 , pp. 503-516
    • Dahanukar, A.1    Lei, Y.T.2    Kwon, J.Y.3    Carlson, J.R.4
  • 35
    • 0018363674 scopus 로고
    • The contribution of a sensitizing pigment to the photosensitivity spectra of fly rhodopsin and metarhodopsin
    • Minke, B. & Kirschfeld, K. The contribution of a sensitizing pigment to the photosensitivity spectra of fly rhodopsin and metarhodopsin. J. Gen. Physiol. 73, 517-540 (1979).
    • (1979) J. Gen. Physiol. , vol.73 , pp. 517-540
    • Minke, B.1    Kirschfeld, K.2
  • 36
    • 0033573379 scopus 로고    scopus 로고
    • Blue-and green-absorbing visual pigments of Drosophila: Ectopic expression and physiological characterization of the R8 photoreceptor cell-specific Rh5 and Rh6 rhodopsins
    • Salcedo, E. et al. Blue-and green-absorbing visual pigments of Drosophila: ectopic expression and physiological characterization of the R8 photoreceptor cell-specific Rh5 and Rh6 rhodopsins. J. Neurosci. 19, 10716-10726 (1999).
    • (1999) J. Neurosci. , vol.19 , pp. 10716-10726
    • Salcedo, E.1
  • 37
    • 84878947585 scopus 로고    scopus 로고
    • Parallel neural pathways mediate CO2 avoidance responses in Drosophila
    • Lin, H.H., Chu, L.A., Fu, T.F., Dickson, B.J. & Chiang, A.S. Parallel neural pathways mediate CO2 avoidance responses in Drosophila. Science 340, 1338-1341 (2013).
    • (2013) Science , vol.340 , pp. 1338-1341
    • Lin, H.H.1    Chu, L.A.2    Fu, T.F.3    Dickson, B.J.4    Chiang, A.S.5
  • 38
    • 49049099862 scopus 로고    scopus 로고
    • A FLEX switch targets Channelrhodopsin-2 to multiple cell types for imaging and long-range circuit mapping
    • Atasoy, D., Aponte, Y., Su, H.H. & Sternson, S.M. A FLEX switch targets Channelrhodopsin-2 to multiple cell types for imaging and long-range circuit mapping. J. Neurosci. 28, 7025-7030 (2008).
    • (2008) J. Neurosci. , vol.28 , pp. 7025-7030
    • Atasoy, D.1    Aponte, Y.2    Su, H.H.3    Sternson, S.M.4
  • 39
    • 84870428606 scopus 로고    scopus 로고
    • Three-dimensional multiwaveguide probe array for light delivery to distributed brain circuits
    • Zorzos, A.N., Scholvin, J., Boyden, E.S. & Fonstad, C.G. Three-dimensional multiwaveguide probe array for light delivery to distributed brain circuits. Opt. Lett. 37, 4841-4843 (2012).
    • (2012) Opt. Lett. , vol.37 , pp. 4841-4843
    • Zorzos, A.N.1    Scholvin, J.2    Boyden, E.S.3    Fonstad, C.G.4
  • 40
    • 84876310253 scopus 로고    scopus 로고
    • Injectable, cellular-scale optoelectronics with applications for wireless optogenetics
    • Kim, T.I. et al. Injectable, cellular-scale optoelectronics with applications for wireless optogenetics. Science 340, 211-216 (2013).
    • (2013) Science , vol.340 , pp. 211-216
    • Kim, T.I.1
  • 41
    • 34247351646 scopus 로고    scopus 로고
    • High Ca2+-phosphate transfection efficiency in low-density neuronal cultures
    • Jiang, M. & Chen, G. High Ca2+-phosphate transfection efficiency in low-density neuronal cultures. Nat. Protoc. 1, 695-700 (2006).
    • (2006) Nat. Protoc. , vol.1 , pp. 695-700
    • Jiang, M.1    Chen, G.2
  • 42
    • 0037593865 scopus 로고    scopus 로고
    • Surface expression of inward rectifier potassium channels is controlled by selective Golgi export
    • Stockklausner, C. & Klocker, N. Surface expression of inward rectifier potassium channels is controlled by selective Golgi export. J. Biol. Chem. 278, 17000-17005 (2003).
    • (2003) J. Biol. Chem. , vol.278 , pp. 17000-17005
    • Stockklausner, C.1    Klocker, N.2
  • 43
    • 0035970773 scopus 로고    scopus 로고
    • A sequence motif responsible for ER export and surface expression of Kir2.0 inward rectifier K+ channels
    • Stockklausner, C., Ludwig, J., Ruppersberg, J.P. & Klocker, N. A sequence motif responsible for ER export and surface expression of Kir2.0 inward rectifier K+ channels. FEBS Lett. 493, 129-133 (2001).
    • (2001) FEBS Lett. , vol.493 , pp. 129-133
    • Stockklausner, C.1    Ludwig, J.2    Ruppersberg, J.P.3    Klocker, N.4
  • 44
    • 57049123832 scopus 로고    scopus 로고
    • ENpHR: A Natronomonas halorhodopsin enhanced for optogenetic applications
    • Gradinaru, V., Thompson, K.R. & Deisseroth, K. eNpHR: a Natronomonas halorhodopsin enhanced for optogenetic applications. Brain Cell Biol. 36, 129-139 (2008).
    • (2008) Brain Cell Biol. , vol.36 , pp. 129-139
    • Gradinaru, V.1    Thompson, K.R.2    Deisseroth, K.3
  • 45
    • 0035846990 scopus 로고    scopus 로고
    • Role of ER export signals in controlling surface potassium channel numbers
    • Ma, D. et al. Role of ER export signals in controlling surface potassium channel numbers. Science 291, 316-319 (2001).
    • (2001) Science , vol.291 , pp. 316-319
    • Ma, D.1
  • 46
    • 78951477322 scopus 로고    scopus 로고
    • Refinement of tools for targeted gene expression in Drosophila
    • Pfeiffer, B.D. et al. Refinement of tools for targeted gene expression in Drosophila. Genetics 186, 735-755 (2010).
    • (2010) Genetics , vol.186 , pp. 735-755
    • Pfeiffer, B.D.1
  • 47
    • 32944482206 scopus 로고    scopus 로고
    • The expression pattern of the Drosophila vesicular glutamate transporter: A marker protein for motoneurons and glutamatergic centers in the brain
    • Mahr, A. & Aberle, H. The expression pattern of the Drosophila vesicular glutamate transporter: a marker protein for motoneurons and glutamatergic centers in the brain. Gene Expr. Patterns 6, 299-309 (2006).
    • (2006) Gene Expr. Patterns , vol.6 , pp. 299-309
    • Mahr, A.1    Aberle, H.2
  • 48
    • 47749087494 scopus 로고    scopus 로고
    • Tools for neuroanatomy and neurogenetics in Drosophila
    • Pfeiffer, B.D. et al. Tools for neuroanatomy and neurogenetics in Drosophila. Proc. Natl. Acad. Sci. USA 105, 9715-9720 (2008).
    • (2008) Proc. Natl. Acad. Sci. USA , vol.105 , pp. 9715-9720
    • Pfeiffer, B.D.1
  • 49
    • 0029814954 scopus 로고    scopus 로고
    • Conditioned visual flight orientation in Drosophila: Dependence on age, practice, and diet
    • Guo, A. et al. Conditioned visual flight orientation in Drosophila: dependence on age, practice, and diet. Learn. Mem. 3, 49-59 (1996).
    • (1996) Learn. Mem. , vol.3 , pp. 49-59
    • Guo, A.1
  • 50
    • 36849028150 scopus 로고    scopus 로고
    • A modular display system for insect behavioral neuroscience
    • Reiser, M.B. & Dickinson, M.H. A modular display system for insect behavioral neuroscience. J. Neurosci. Methods 167, 127-139 (2008).
    • (2008) J. Neurosci. Methods , vol.167 , pp. 127-139
    • Reiser, M.B.1    Dickinson, M.H.2


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