培養神経回路網の長期的観測による
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- しなつ ゆきしげ
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1 ( )
2 64 0.1ml/h 5DIVDays in vitro 50msec k-means 2
3 Amp width i
4 ANTTmain...72 B mlt...73 ii
5 iii G0D DIV 12DIV DIV 20DIV DIV 51DIV
6 iv
7 [1][2][3] [4] in vivo 1979 Gross
8 2 [5][6] (synchronized bursting) [7] [8][9] days in vitro (DIV) 50 5DIV 13DIV 15DIV
9
10 ATP Na K -60mV NaNa K K Na Na +40mV K K 1msec Hodgikin-Huxley [10] Na 1msec Hodgikin-Huxley [11] m/sec /sec 1msec EPSPExcitatory Post-Synaptic Potential GABA-aminobutyric acid GABA IPSPInhibitory Post-Synaptic Potential EPSP IPSP non-nmda (N-methyl-D-aspartate) NMDA non-nmda (AMPA ) Na + K + 4
11 non-nmda EPSP NMDA Mg 2+ non-nmda EPSP NMDA non-nmda EPSC [12] NMDA NR1 4 (NR2A-NR2D) [13] GABA [14] GABA muscimol 5
12 2-1 Chemical Abbrev. Transmitter blocked Picrotoxin GABA Bicuculline GABA 6-cyano-7-nitroquinoxaline-2,3-dione CNQX Glutamate - Kainate 6-cyano-7-nitroquinoxaline-2,3-dione CNQX Glutamate - AMPA D-2-amino-5-phosphonovalerate APV or AP5 Glutamate - NMDA Magnesium++ Mg++ Glutamate - NMDA D(-)-2-amino-7-phosphono-heptanoate A7 Glutamate - NMDA 3-((+/-)-2-carboxypiperazin-4-yl)propyl-1-phosphonic acid CPP Glutamate - NMDA Strychnine Glycine D-tubocurarine ACh curare occupies ACh receptors tetrodotoxin TTX sodium channels saxitoxin STX sodium channels Procaine sodium channels Tetraethylammonium ion TEA potassium channels cholera toxin A subunit CTX GTP -> GDP pertussis toxin A subunit PTX GDP -> GTP SKF81297 Dopamine (D1) Quinpirole Dopamine (D2) spiperone Dopamine (D2) Clozapine Dopamine (D4?) 6
13 LGNlateral ganiculate nucleus 1991 M. Meister [4] in vivo [15] in vitro [16] NMDA Mg 2+ Ca 2+ H. P. C. Robinson [17] NMDA APV [18] 30 [19] [20]
14 H. Kamioka [7] mm/s sec cm/s 40sec 30 (450msec) (100msec) APV, TTX, muscimol NMDA 30 APV NMDA NMDA TTX CNQX APV non-nmda Hebb
15 9 2.3 S/N EPSPIPSP S/N 1979 Guenter W. Gross 2 S/N 100V1msec V 300Hz6Hz
16 [23] [24][25] Mg 2+ 50DIV
17 m 180m ml 25Hz16bit DIV A B 150msec B1ch 11
18 500m
19 incubator acrylic cap 64 microelectrode array
20 A 51 DIV B1ch D0ch E0ch F0ch G0ch H0ch 100V -100V B 150sec B1ch D0ch E0ch F0ch G0ch H0ch 100V -100V 150msec
21
22 3.2.1 Amp x n ( n = 1,2,..., n) x n < threshold 3-1 xn x n 3-6 Baseline = 1 Spike_ size 1 Spike_ size 2 X Po + n n 1 Valley _ = Spike_ size int Amp = X Baseline 3-3 Valley_ Point Spike-size Amp Base-line
23 3.2.2 width 3-7 t t 2 t1 = 1/ f[ Hz] 3-4 a = threshold V ( t ) b = V( t2) threshold t i b = t ( ) 2 t 2 t1 a + b t o i 3-6 t 4 t3 = 1/ f [ Hz] 3-7 c = threshold V ( t ) d = V ( t3) threshold t o = t t 3 ( t4 3 ) d c + d 3-9 width = t o t i
24
25
26 C shape ( a, b) = + i + τ τ i= τ = τ S ( i) S ( i) 2 a a S ( i) b + τ i= τ S ( i) 2 b 3-11 { Sa (0), Sb (0)} { S (0), S (0)} min C amp ( a, b) = 3-12 max a b τ = ( S i τ a ( i) 2 d ( a, b) = S ( i)) 3-13 b 20
27 3.3.3 num a num b 2 g( t i a i t late t = = b j = 1 j 1,,, ) C ( a, b, t late ) num + num t1 a g( t1, t 2) = b t2 < τ 1 else
28 3.3.4 late 1, late 2, L, late N N num e g t t late e i e i e f t = = (, 1 1, ) ( ) = 3-15 N 22
29 V C3H2 H3 E E0 1msec 1msec 2 2 E0 A B A D0,E0,E1,F1,G0,H0 B B0,B1,D0,E0,E A B B1ch B A B 3-13 / AB 50DIV 80DIV 3-14 A 1msec 10msec AB 50DIV 54DIV, 57DIV A 50DIV 2.5msec 61DIV 2.7msec 82DIV 5.6msec 23
30 51 DIV 80 [V] [msec] 24
![60[V] 0.1 51 DIV E0 ch 0 0.](/docs-images/93/113580891/images/31-1.jpg "4 [msec] 0 20[msec] 60[V] E0 ch 0.1 A 0 0.")
31 60[V] DIV E0 ch [msec] 0 20[msec] 60[V] E0 ch 0.1 A [msec] B 0 20[msec]
32 A B C D ( [msec]) 25msec -100V 100V B0ch B1ch D0ch E0ch E1ch F1ch G0ch H0ch A A A B B
33 DIV 50 A Log-scale B Log-scale ms5ms10ms 1ms 10ms 100ms 1s1ms 10ms 100ms 1s
34 DIV 82DIV 4
35 in vitro AD ph CO2 DIV
36 Wistar 0.02 (Boehringer) 3620 D DMEMDulbecco s Modified Eagle s Medium: Gibico 5% Gibco5%2.5g/ml (Sigma) / 5?40U/ml 1 conditioned medium ITOIndium-tin-oxide m 2 30m 180m
37 500m 180m
38 in vitro CO 64 (10ml) 0.1ml/h CO 10%37
39 4-4 33
40 4-5 34
41 V Hz 64 25kHz 16bit 276GB CO 4-6
42 4-6 36
43 A/D (500MB) 30 25kHz 16bit MB msec 1msec MB 5DIV 10kB16DIV 10MB [mlt ] B
44 4-8 38
45 CO2 OFF/ON CO OFF/ON 4-9 CLOSE OPEN 30V OPEN 5msec msec
46 4-9 40
47 ) ( ) (,, a e a e t t t I = δ 4-1 h(t) > = 0) ( 0 2) / ( 1/ ) ( 0 t T t t h τ 4-2 e a u e,a (t)
48 u τ / 2 ( t) = h( t τ ) I e, a( t) dτ = h( t te, e, a a τ / 2 ) num e r ( t) = u ( t e i= 1 e, i ) R( t) = 64 e= 1 r e ( t) h( t) = 1 2πT 2 0 e 2 t 2T
49 D( a, b) = ( Z a, e Zb, e ) e= Z n e Sn, e Se = σ, S e 43
50
51
52 DIV 51DIV DIV 51DIV 54DIV DIV DIV DIV 4-12
53 DIV 53DIV G0 D3 1.6 V G0 D3 19DIV 4-14 G0 21DIV D3 425DIV 52DIV D DIV 42 5DIV 47
54 G0D3 A0 B0 C0 D0 E0 F0 G0 H0 A1 B1 C1 D1 E1 F1 G1 H1 A2 B2 C2 D2 E2 F2 G2 H2 A3 B3 C3 D3 E3 F3 G3 H3 A4 B4 C4 D4 E4 F4 G4 H4 A5 B5 C5 D5 E5 F5 G5 H5 A6 B6 C6 D6 E6 F6 G6 H6 A7 B7 C7 D7 E7 F7 G7 H7 DIV V DIV V
55 4.3.2 DIV 53DIV DIV B3,C0,D4,E7,F7,F4,H3,DIV B3 14DIV15DIV 16DIV H3,A6 49DIV 50DIV 51DIV DIV 19DIV 13DIV 19DIV 49
56 5 DIV 6 DIV 100V 7 DIV 8DIV 150sec 9 DIV 10 DIV 11 DIV 12 DIV 4-155DIV 12DIV 50
57 13 DIV 14 DIV 100V 15 DIV 16 DIV 150sec 17 DIV 18 DIV 19 DIV 20 DIV DIV 20DIV 51
58 23 DIV 27 DIV 100V 32 DIV 37 DIV 150sec 43 DIV 49 DIV 50 DIV 51 DIV DIV 51DIV 52
59 A 13DIV B 19DIV 150sec
60 4.3.3 DIV 50DIV 1sec DIV 4-21 log-scale 4-20 DIV 1 /sec 13.5DIV 10 /sec 2 15DIV 0/sec 15DIV 25DIV 300 /sec 15DIV 13.5DIV DIV A1, B4, D4 13.5DIV A0,A5,F7 1 H3 13.5DIV H DIV 4-20-A 600 /sec 700 /sec 4-20-B log-scale 20DIV 28DIV DIV 15DIV log-scale 15DIV 10sec 100sec 10DIV 1sec 10sec 4-18-A A) B 15DIV 19DIV 3sec 25DIV 2sec 15DIV 18DIV 15DIV 54
61 DIV A B A DIV 13DIV 15DIV 15DIV 5 B 15DIV DIV DIV 15DIV 1sec 150sec 55
62 (5-50DIV) A0 B0 C0 D0 E0 F0 G0 H0 A1 B1 C1 D1 E1 F1 G1 H1 A2 B2 C2 D2 E2 F2 G2 H2 A3 B3 C3 D3 E3 F3 G3 H3 A4 B4 C4 D4 E4 F4 G4 H4 A5 B5 C5 D5 E5 F5 G5 H5 A6 B6 C6 D6 E6 F6 G6 H6 A7 B7 C7 D7 E7 F7 G7 H7 DIV /sec
63 (9-16DIV) A0 B0 C0 D0 E0 F0 G0 H0 A1 B1 C1 D1 E1 F1 G1 H1 A2 B2 C2 D2 E2 F2 G2 H2 A3 B3 C3 D3 E3 F3 G3 H3 A4 B4 C4 D4 E4 F4 G4 H4 A5 B5 C5 D5 E5 F5 G5 H5 A6 B6 C6 D6 E6 F6 G6 H6 A7 B7 C7 D7 E7 F7 G7 H7 /sec DIV
64 (/sec) A DIV (/sec) Blog DIV
65 DIV
66
67 DIV 15DIV DIV 25DIV A DIV A B 5A A B A 15DIV 3 C H3ch D C D E C D E 15DIV A 12.5DIV 15DIV 16DIV C,D,E DIV A,B C,D,E A 14DIV DIV H3ch E4ch 4-25 A H3, E4ch 14.5DIV A 4.0 A H3ch H3,E4ch C 61
68 DIV CRT 4-27
69
70 14.5DIV A5 B3 C0 E0 C3 A6 E4 H3 5.5hour 3.5hour 4.0hour 100V -100V 150sec
71 500m 180m
72 CO2 2 3DIV 30DIV 5DIV 5DIV 13DIV 13DIV 15DIV 15DIV 15DIV 15DIV 15DIV 15DIV 4-19 H3 H DIV
73 DIV NMDA DIV 50DIV 2
74
75 68 NTT
76 [1] T. Tsumoto : Long-term potentiation and log-term depression in the neocortex, Progress in Neurobiology : Vol.39, pp , (1992) [2] Y. Jimbo, T. Tateno, H. P. C. Robinson : Simultaneous induction of pathway-specific potentiation and depression in Networks of cortical neurons : Biophysical Journal, 76, pp , (1999) [3] D. Debanne, B. H. Gahwiler, S. M. Thompson : Long-term synaptic plasticity between pairs of individual CA3 pyramidal cells in rat hippocampal slice cultures : J. Physiol., 507.1, pp , (1998) [4] M. Meister, R. O. L. Wong, D. A. Baylor, C. J. Shatz : Synchronized Bursts of Action Potentials in Ganglion Cells of the Developing Mammalian Retina : Science, 252, pp , (1991) [5] G.W.Gross, Simultaneous Signals Unit Recording in vitro with a Photoetched Laser Deinsulated Gold Multimicroelectrode Surface : IEEE Trans. Biomed. Eng., 26, pp , (1979) [6] : : : Vol.6, No.1, pp.24-29, (1999) [7] H. Kamioka, E. Maeda, Y. Jimbo, H. P. C. Robinson, A. Kawana : Spontaneous periodic synchronized bursting during formation of mature patterns of connections in cortical cultures : Neurosci. Lett., 206, pp , (1996) [8] G.W.Gross, F.U.Schwalm : A closed flow chamber for long-term multichannel recording and optical monitoring : J. Neurosci. Methods, 52, pp.73-85, (1994) [9] S. M. Potter, T. B. DeMarse : A new approach to neural cell culture for long-term studies : J. Neurosci. Methods, 110,pp.17-24,(2001) [10] A. L. Hodgkin, B. Katz : The effect of sodium ions on the electrical activity of the giant axon of the squid : J. Physiol., 108, 37-77, (1949) [11] : :, (1995) [12],, : :, pp , (1996) [13] Ratna Sircar : Developmental maturation of the N-methl-D-aspartic acid receptor channel complex in postnatal rat brain : Devl. Neurosci., 18, pp , (2000) [14] S. Watanabe, Y. Jimbo, H. Kamioka, Y. Kirino, A. Kawana : Developmental of low 69
77 magnesium-induced spontaneous synchronized bursting and GABAergic modulation in cultured rat neocortical neurons : Neurosci. Lett., 210, pp.41-44, (1996) [15] M. Steriade, F. Amzica, A. Nunez : Cholinergic and Noradrenergic Modulation of the Slow Oscillation in Neocortical Cells : J. Neurophysiol., 70, pp , (1993) [16] Y. Jimbo, H. P. C. Robinson : Propagation of spontaneous synchronized activity in cortical slice cultures recorded by planar electrode arrays : Bioelectrochem.,51, pp ,(2000) [17] H. P. C. Robinson, M. Kawana, Y. Jimbo, K. Torimitsu : Periodic Synchronized Bursting and Intracellular Calcium Transients Elicited by Low Magnesium in Cultured Cortical Neurons : J. Neurophysiol., 70, pp , (1993) [18] K. Nakanishi, F. Kukita : Functional synapses in synchronized bursting of neocortical neurons in culture : Brain. Res., 795, pp , (1998) [19] E. Maeda, H. P. Robinson, A. Kawana : The mechanisms of generation and propagation of synchronized bursting in developing networks of cortical neurons : J. Neurosci., 15, pp , (1995) [20] R.Segev, Y.Shapira, M.Benveniste, E.Ben-Jacob : Observations and modeling of synchronized bursting in two-dimensional neural networks, Physical Review E, Vol.64, [21] Y. Jimbo, A. Kawana, P. Parodi, V. Torre : The dynamics of a neuronal culture of dissociated cortical neurons of neonatal rats : Biol. Cybern., 83, pp.1-20, (2000) [22] O. Klostermann, P. Wahle : Patterns of spontaneous activity and morphology of interneuron types in organotypic cortex and thalamus-cortex cultures : Neurosci. Vol. 92, pp , (1999) [23] N. Konno, T. Fukami, T. Shiina, Y. Jimbo : Estimation of network structure for signal propagations by the analysis of multichannel action potentials in cultured neural networks : Trans. IEE of Japan, 188, pp , (1998) [24] G. Zouridakis, D. C. Tam : Multi-unit spike Discrimination using wavelet transforms : Comput. Biol. Med., Vol.27, No.1, pp.9-18, (1997) [25] E. Hulata, R. Segev, Y. Shapira, M. Benveniste, E. B. Jacob : Detection and sorting of neural spikes using wavelet packets : Physical review Lett., 85, pp , (2000) [26] D. A. Stenger, T. M. McKenna : Enabling Technologies for Cultured Neural Networks : Academic Press, INC., (1994) [27] H. Eichenbaum, J. L. Davis : neural ENSEMBLES : Wiley-Liss, (1998) [28],, : :, 152, pp15-20, (2000) [29],, : :, pp.31-34, (2001) [30],, : 0 ME, (2001) [31],, : 70
78 : C [32] : :, (1996) [33] : :, (1997) [34] : :, (1995) [35] : : 8 C, (1997) 71
79 72 ANTTmain
80 B mlt 0 1(char ) 100 multidata-packet-ver (long ) 4(int ) 4(int ) 4(int ) 4(int ) 4(int ) 4(int ) (pt) 20k 0 1 (pt) 128 4(int ) 64 (A0,A1,,A7,B0,B1,,H7) 384 4(int ) 64 (pt) 4(int ) (pt) 2(short int ) 64 4(float ) 64 4(float ) (int ) 64 4(int ) 4(float ) 4(int ) 64 (pt) (pt) 73
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