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TRANSCRIPT
2012
S2 Mn
G
261102010
物性研究・電子版 Vol, 2, No. 2, 022601 (2013年5月号)《修士論文》
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II PS II Mn
Mn 4 Mn 5 O 1 Ca
Mn 5 S0, S1, S2,
S3, S4 S1 X S2
Mn III, IV, IV, IV
ENDOR Mn
PELDOR
ESR Mn S2
S2 Mn PS II YD• PELDOR
PELDOR YD•-S2 Mn
R = 27 Å YD•-Mn YD
30 Å YD• Mn
spin projection PELDOR
PSII PELDOR
spin projection
Mn S2
4 Mn Mn1, 2, 3, 4 Spin projection [1.97, -1.20,
1.19, -0.96] Mn [III, IV, IV, IV]
spin projection Mn1-3 J13
Mn
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1
1.1 ..................................................................................................................................... 4
1.2 II .............................................................................................................................. 5
1.3 Mn ....................................................................................................................... 7
1.4 Mn ........................................................................................... 8
1.5 Mn S2 ...................................................................................................... 8
1.6 Spin projection ......................................................................................................................... 9
1.7 Mn S2 ......................................................... 10
1.8 ....................................................................................................................... 12
2
2.1 ....................................................................................................................................... 13
2.1.1 PS II ........................................... 13
2.1.2 T. vulcanus PS II ................................................... 14
2.1.3 S1 100% PS II .................................................................................... 15
2.1.4 S2 PS II .................................................................................. 16
2.1.5 PS II ....................................................................................................... 17
2.2 ............................................................................................................................... 18
2.2.1 ESR ......................................................................................... 18
2.2.2 ESR ............................................................................................................... 21
2.2.3 PELDOR ................................................................. 22
2.3 PELDOR ................................................................................................... 23
3
3.1 ESR ................................................................................................... 25
3.1.1 S2 Mn ESE ................................................ 25
3.1.2 YD• CW-ESR ........................................................................ 26
3.2 PS II S2 Mn -YD• PELDOR ........................................ 27
3.3 PSII S2 Mn -YD• PELDOR ............................................. 32
4 .............................................................................................................................................. 36
5 .......................................................................................................................................... 37
6 ...................................................................................................................................... 38
7 .............................................................................................................................................. 40
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1 1.1
ATP
2
NADPH
4 1-1 4
II PS II I PS I b6f ATP
PS I PS II
PSII
PS II P680 P680
PS II
QB QB 2 [McEvoy and
Brudvig 2006; Nelson and Yocum 2006; Messinger and Renger 2008] 1.2
b6f PC PS I
P700 P700 P680
1-1
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PS I FD
-NADP FNR NADP+
NADPH NADPH
ATP
2 ADP 3 ATP ATP
NADPH
1.2 II
PSII PS II D1/D2
CP43/47 1-2 PS II
LHC(Light harvesting complex)II CP43/47
LHC II P680 D1/D2 C2
2
PSII
PS II PsbO, PsbU, PsbV PS II
PsbO, PsbP, PsbQ [McEvoy and Brudvig 2006; Nelson and Yocum 2006;
Messinger and Renger 2008]
1-2 T. vulcanus PS II
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D1/D2 D/Z YD/YZ P680
2 a Chl a Pheo QA/QB
Fe
P680 D1
D2 QA QB P680
D1 YZ
YZ Mn Mn
D1/D2
D2 D1 YZ YD
YD Mn
Mn
1-3 PS II
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1.3 Mn
Mn Mn
(Oxygen evolving complex)
1-4 X 1.9 Å
[Umena et al. 2011] Mn 4 Mn Ca Cl
PS II Mn Ca Cl
4 Mn Mn1 Mn2 Mn3 Mn4 Ca
Mn1 His332 D342 E189 Mn2
D342 A344 D2-E354 Mn3 E333 D2-E354 Mn4 E333 D170
1-4 X PS II
[Umena et al. 2011]
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1.4 Mn
Mn 5 S0-4 [Kok et al. 1970]
1-5 S0 1 Mn 1
S3 Mn
S4 S0 2 1
S1 S4
S0
1.5 Mn S2
S2 Mn 1 III 3 IV Mn
1-5 III IV Mn S = 2, 3/2
Mn Mn
ST = 1/2 ESR
S2 Mn ESR g = 2
[Dismukes and Siderer 1981]
1-5 Mn
S 4 Mn
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1.6 Spin projection
Mn
i Mn (ZFS)
g a i j
ZFS 1
i ZFS
3
1-6 Spin projection ST Si
i Mn Spin projection ST Siprojection
Si ST >90
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1.7 Mn S2
ESR Mn
S2 Mn Mn J
Mn X-, Q- 55Mn-Pulsed ENDOR
Mn(III) -165 ~ -225 MHz Mn(IV) -187 ~ -253MHz
[Peloquin et al. 2000; Charlot et al. 2005; Kulik et al. 2005; Cox et al. 2011] Mn
spin projection 1
1 Pulsed ENDOR Mn spin projection
Mn(III) Mn(IV) Mn(IV) Mn(IV)
1.70 1.27 -1.0 -0.99 [Peloquin et al. 2000]
1.73 1.10 -1.04 -0.79 [Kulik et al. 2005]
1.73 1.12 -0.95 -0.90 [Cox et al. 2011] a1.39 ~ 1.89 a 0.99 ~ 1.34 a -1.11 ~ -0.82 a -1.02~ -0.75 [Cox et al. 2011] a Mn(III) Mn(IV) Aiso 165 ~ 225 187 ~ 253 MHz
Mn J
1-7 Peloquin Mn S2 4
[Peloquin et al. 2000] Peloquin 4 Mn
Kulik 3 Mn MnB(IV), MnC(III), MnD(IV)
MnB MnA Y [Kulik et al. 2005] Siegbahn
EXAFS Mn MnB(IV) 4
Mn [Pantazis et al. 2009] Cox Q 55Mn-ENDOR DFT Siegbahn [Siegbahn 2008;
Siegbahn 2009; Siegbahn 2009; Cox et al. 2011]
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ESEEM ENDOR
ESEEM Mn(III) 14N
[Yeagle et al. 2008; Stich et al. 2011] X
Mn N His332
Glu Mn(III) His332
1-7 S2 Mn
1) Peloquin 2) Kulik DFT
3) Pantazis EXAFS
4) Siegbahn Pantazis DFT
[cm-1]
1-3 MnA-B 4 2-4 MnC-D
3 4 MnA
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PELDOR
2.2.4 S2 Mn YD PELDOR
27 Å [Hara et al. 1996] 1-8
X YD Mn 29 - 32 Å
PELDOR Mn 4 Mn
Spin projection
1.8
Mn S2 Mn Spin
projection YD-S2 Mn
PELDOR Mn S2 X
Mn Spin projection Mn
1-8 YD-S2 Mn PELDOR
27 Å 30 Å
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2 2.1
2.1.1 PS II
BBY PS II
[Ono and Inoue 1986; Banham et al. 2008] 0
PSII
(1) A
0.4 M sucrose, 15 mM NaCl, 2 mM EDTA, 40 mM Hepes/NaOH (pH=7.5), 3 %BSA
(2) B
0.4 M sucrose, 15 mM NaCl, 5 mM MgCl, 40 mM Mes/NaOH (pH=6.5)
(3) C
0.4 M sucrose, 15 mM NaCl, 40 mM Mes/NaOH (pH=6.5), 30 % ethylene glycol
A
↓
5,500 ×g R9A 7,000 rpm 25
↓
B
↓
100 ×g R20A2 1,000 rpm 4
↓
↓
30,000 ×g R20A2 17,000 rpm 5
↓
B [Chl] = 5 mgChl/ml
[Chl] = 3 mgChl/ml 6.3 % Triton-X B
20
↓
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32,600 ×g R20A2 18,000 rpm 25
↓
B
↓
7,300 ×g R20A2 8,500 rpm 15
↓
↓
32,600 ×g R20A2 18,000 rpm 25
↓
B
↓
32,600 ×g R20A2 18,000 rpm 25
↓
PSII
C 5 mgChl/ml
2.1.2 T. vulcanus PS II
T. vulcanus PS II [Shen and Inoue 1993;
Shen and Kamiya 2000] T. vulcanus PS II
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2.1.3 S1 100% PS II
PS II Mn
S1 PS II 273
S0-S3 25% PS II
S0 S2 S3 S1 S0:S1 25:75 S0
YD• S1 S0YD
• S1YD [Styring and Rutherford
1987]
S1 100 PS II
S1 100% PS II
D
0.4 M Sucrose, 20 mM NaCl, 1 mM EDTA, 50 mM Mes/NaOH (pH = 6.5)
BBY PS II
D buffer [Chl] = 1 mgChl/ml
273 K 1 2
S1 100% PS II
32,600 ×g R20A2 18,000 rpm 20
ESR
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2.1.4 S2 PS II
200 K S2 S1 250 K
QA QB S2 S3
200 K S2 PS II
ESR PS II
200 K
S1 100% PS II 200 K
LED 150 μmol m-2 s-1
5
ESR
2-1 ESR
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2.1.5 PS II
BBY PS II
PS II
PS II
PS II OHP
4 15
OHP 2.5 mm 25 mm 6
ESR
S1 S2 PS II
2-2
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2.2
2.2.1 ESR
ESR
1 μ
(6)
g
(7)
e
2 , 2
(8)
α, β
2 Zeeman
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2-3
ESR 2-3
CW-ESR
1 1
H
g
Zeeman Zeeman
2
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R
A B
C, D
E, F
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2.2.2 ESR
CW-ESR
ESR tp
ns
2-4 z
(a)
x tp z y
xy (b) xy xy
(c) 90° τ tp 180°
yz xy (d)
τ y (e)
2-4 ESE
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2.2.3 PELDOR
PELDOR 2
2 1,2
15 A, B
1, 2 1, 2 g 2
H
2-5 PELDOR ν1,2 1,2
1 ν1 90° (1st) 180°
(3rd) ν2 180° (2nd) 2
1 2 1st
2nd T 1 ESE
p 2nd
2-5 PELDOR
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2.3 PELDOR
PS II YD•-Mn YD
•
6 C O Mn 4 Mn
spin projection
1 YD•-Mn DT 17
YD• i (i = 1-7) Mn j (j = 1-4)
spin projection YD•-Mn
D
PELDOR 18
PS II C2 z
PS II PELDOR
Gauss PS II
H 0 90
2-6 PS II
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X 1.9 Å
PELDOR [Umena et al. 2011] YZ YD
Fe PS II C2 Tyr
ENDOR 2-7 4 Mn spin projection 1
[Mino et al. 1997; Nagai et al. 2008; Kessen et al. 2010]
2-7 YD•
[Nagai et al. 2008]
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3 3.1 ESR
3.1.1 S2 Mn ESE
3-1 T. vulcanus PS II A S2 B S1 ESE
g = 2 60 mT ST = 1/2 S2 Mn
S2 YD•
YD• PS II
T. vulcanus
PELDOR S2
YD•
3-1 T. vulcanus PS II ESE
(A)S2 (B)S1
9.60 GHz
τ 200 ns
2 ms
5 K
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3.1.2 YD• CW-ESR
3-2 YD• g = 2 CW-ESR
T. vulcanus PS II S1 S2 S2 340 mT
3 mT ChlZ+ ChlZ
+ S2
ChlZ+ 338.5 mT
ChlZ+ PS II 8% T. vulcanus PS II 15%
ChlZ Mn YD ChlZ-S2 Mn
PELDOR YD ChlZ+
PELDOR
3-2 YD
• CW-ESR
9.592 GHz
50.5 μW
4 G
170 K
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3.2 PS II S2 Mn -YD• PELDOR
3-3 PS II S2 Mn -YD• PELDOR
T. vulcanus A B PS II
PS II A,
B PS II T. vulcanus
3.2.1
PELDOR S2 Mn -YD• 27 Å
X Mn-YD• YD
Mn1-4 29.1 31.9 31.1 30.1 Å 3-3
27 C 30 Å D Mn-YD•
Mn 4 Mn Spin
projection
YD• Mn
3-3 S2 Mn - YD•
(A)T. vulcanus (B) PELDOR
(C)27 Å (D)30 Å
ν1 = 9.61 GHz ν2 = 9.73 GHz τ = 1000 ns
348 mT 2 ms 5 K
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3.2.2
YD• S2 Mn 4 Mn
S2 Mn spin projection ENDOR DFT
1.7 Mn spin projection Mn
spin projection Peloquin Mn spin projection
1.7, 1.27, -1, -0.99 [Peloquin et al. 2000] -1 -0.99 -1
Mn spin projection 12
Peloquin spin projection 3-4 Mn
spin projection 2
3-3 A b2 c2 d2 Mn1
spin projection 1.7 a1-3 Mn4 1.7 Mn1 1.27
d1 spin
projection
2 3-4 Mn spin projection
Mn1 Mn2 Mn3 Mn4
a1 1.7 1.27 -1 -1
a2 1.7 -1 1.27 -1
a3 1.7 -1 -1 1.27
b1 1.27 1.7 -1 -1
b2 -1 1.7 1.27 -1
b3 -1 1.7 -1 1.27
c1 1.27 -1 1.7 -1
c2 -1 1.27 1.7 -1
c3 -1 -1 1.7 1.27
d1 1.27 -1 -1 1.7
d2 -1 1.27 -1 1.7
d3 -1 -1 1.27 1.7
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3-4 S2 Mn - YD• PELDOR
Mn Spin projection 1.7 1.27 -1.0 -1 [Peloquin et al. 2000]
Spin projection 1.7 Mn(III) a1-3 Mn1 b1-3 Mn2 c1-3 Mn3
d1-3 Mn4 2 Spin projection 1.27 Mn a1-3
Mn2-4 b1-3 Mn1,3,4 c1-3 Mn1,2,4 d1-3 Mn1-3
3-2 τʹ = 440, 800 ns
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Mn(III) Mn(IV) Aiso 165 ~
225 187 ~ 253 MHz Cox Mn spin projection
[1.39 ~ 1.89, 0.99 ~ 1.34, -1.11 ~ -0.82, -1.02 ~ -0.75] [Cox et al.
2011] 1 10% [1.2 ~ 2.0, 0.9 ~ 1.5, -1.2 ~ -0.65, -1.2 ~
-0.65] Mn spin projection 12 4
spin projection 1.00 0.01
3-5 B, C, D τʹ = 200, 440, 800 ns
3-4 a2 a3 d1 5% 3
spin projection [++] spin projection [+]
spin projection [-] 3-5 B, C, D Mn spin projection
Mn1, 2, 3, 4 = [++, -, +, -], [++, -, -, +], [+, -, -, ++]
3-5 spin projection
Mn1 spin projection Mn2 Mn3 4
Mn1-2 Mn3-4
3 spin projection
Mn1 Mn2 Mn3 Mn4 Combinations Error range
1.68 ~ 2.00 -1.20 ~- 0.65 0.90 ~1.50 -1.20 ~ -0.65 [++, -, +, -] < 5%
1.20 ~ 1.71 -1.09 ~ -0.65 -1.20 ~ -0.65 0.90 ~ 1.50 [++, -, -, +] < 5%
0.90 ~ 1.50 -1.19 ~- 0.65 -1.20 ~ -0.65 1.20 ~ 2.00 [+, -, -, ++] < 5%
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3-5 PELDOR
A: 3-3 A B-D:
Mn Mn1-4 spin projection
B: 1.77, -1.07, 1.09, -0.75
C: 1.39, -0.75, -1.05, 1.01
D: 1.17, -0.77, -0.77, 1.39
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3.3 PSII S2 Mn -YD• PELDOR
3.2 Mn spin projection Mn
spin projection [++, -, +, -], [++, -, -, +], [+, -, -, ++]
3 PS II
PELDOR
3-6 PS II Cytb559 Cytb559 gZ
n H0 θ = 90° Gauss
Δ = 17 ± 0.5°
3-6 Cytb559 CW-ESR
Cytb559 gZ CW-ESR
9.44 GHz
201 μW
5 K
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3-7 PS II θ = 0° A θ = 90° B YD•-Mn PELDOR
θ = 0° R H0 1
1 θ = 0° τʹ = 220 620 ns θ = 90°
τʹ = 180 580 ns θ = 90° τʹ = 300 ns θ = 0°
τʹ = 430 810 ns θ = 90°
θ = 0° τʹ = 220 430 810 ns θ = 90° τʹ = 180 430 810 ns
3-7 A B Mn1
Mn3 spin projection [++, -, +, -]
Mn1-4 spin projection [1.97, -1.20, 1.19, -0.96]
3-7 S2 Mn - YD• PELDOR
A: H0 n θ = 0° B: θ = 90°
[++, -, +, -] [++, -, -, +] [+, -, -, ++]
ν1 = 9.61 GHz ν2 = 9.73 GHz τ = 1000 ns
348 mT 2 ms 5 K
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spin projection 4
5% [++, -, +, -] 5%
5% spin projection
[++, -, +, -] spin projection [++] 1.87-2.00
4 spin projection
Mn1 Mn2 Mn3 Mn4 Combinations Error range
1.87~ 2.00 -1.20~ -1.03 0.90 ~ 1.25 -1.05 ~ -0.65 [++, -, +, -] < 5%
1.38 ~ 1.41 -0.66 ~ -0.65 -0.67 ~ -0.65 0.90 ~ 0.94 [++, -, -, +] > 5%
1.06 ~ 1.25 -0.65 ~ -0.80 -0.92 ~ -0.65 1.21 ~ 1.50 [+, -, -, ++] > 5%
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PELDOR τʹ = 180
3-8 spin projection
τʹ = 175 - 185 ns 5% [++, -, +, -] [++, -,
-, +] [+, -, -, ++] A B spin projection [++] [+]
C D spin projection [-]
C Mn2 D spin projection [-] [++, -, +, -] Mn4 [++, -, -, +]
[+, -, -, ++] Mn3 [++, -, -, +] [+, -, -, ++] 5%
[++] 1.38-1.41
1.21-1.5 4 3-8A 3-7 [++,
-, -, +] [+, -, -, ++]
3-8 spin projection PELDOR
A spin projection B spin projection
C Mn2 spin projection
D Mn3 ([++, -, -, +], [+, -, -, ++]), Mn4 ([++, -, +, -]) spin projection
物性研究・電子版 Vol, 2, No. 2, 022601 (2013年5月号)《修士論文》
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4 Mn ESR S2 Mn
55Mn-ENDOR S2 4 Mn (III, IV, IV, IV)
PELDOR [++, -, +, -]
[++, -, +, -] spin projection 1.87-2.0
[++, -, -, +] 1.38-1.41 [+, -, -, ++] 1.21-1.5
14N ESEEM His332 Mn Mn(III)
His332 Mn Mn1 PELDOR [++, -, +, -] [++, -, -, +]
ESEEM Mn(III) spin projection 2
[++, -, +, -] 2 spin projection
ESEEM
[++, -, +, -] Mn J 4-1
Mn1-2 Mn3-4 X
Mn1-2 Mn2-3 2.8-2.9 Å Mn
4-1 X
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DFT Mn1,4 spin projection [++, -, -, +] S2
Mn [Ames et al. 2011; Cox et al. 2011]
PELDOR [++, -, +, -]
2
(1) PELDOR Mn
(2) Spin projection
(1) PELDOR Mn
X PS II S1 S1
S2 Mn 0.3 Å Ames
DFT [Ames et al. 2011] PELDOR X
Mn 0.3 Å PELDOR
[++, -, +, -] [++, -, -, +] 5% [+, -, -, ++]
10%
Mn [++, -, +, -] Mn
PELDOR
(2) Spin projection
Ames 55Mn-ENDOR DFT Mn
DFT Mn
PELDOR
Mn spin projection
PELDOR DFT
Mn
DFT [++, -,
-, +]
5 S2 Mn -YD
• PELDOR X
spin projection
Mn1 Mn3 spin projection Mn1-4 spin projection [1.97, -1.19,
1.17, -0.95] Mn1 Mn S2 Mn(III)
Mn1-2 Mn3-4 Mn1-3
[Asada et al. 2013]
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6 Ames W., Pantazis D. A., Krewald V., Cox N., Messinger J., Lubitz W. and Neese F., Theoretical
Evaluation of Structural Models of the S2 State in the Oxygen Evolving Complex of Photosystem II: Protonation States and Magnetic Interactions, Journal of the American Chemical Society, 133 (2011) 19743-19757.
Asada M., Nagashima H., Koua F. H. M., Shen J-R., Kawamori A. and Mino H., Electronic Structure of
S2 state of the oxygen-evolving complex of Photosystem II studied by PELDOR, Biochimica et Biophysica Acta, 1827 (2013) 438-445.
Banham J. E., Baker C. M., Ceola S., Day I. J., Grant G. H., Groenen E. J. J., Rodgers C. T., Jeschke G.
and Timmel C. R., Distance measurements in the borderline region of applicability of CW EPR and DEER: A model study on a homologous series of spin-labelled peptides, Journal of Magnetic Resonance, 191 (2008) 202-218.
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