2012 º Ø 1= emercury.yukawa.kyoto-u.ac.jp/~bussei.kenkyu/pdf/02/2/...2.1.2 4ß(ò$Î#Õ q ö b 6...

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》

2

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


3

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


4

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


5

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


6

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


7

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]


8

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


9

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


10

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]


11

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


12

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


13

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

↓


14

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


15

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


16

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


17

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


18

2.2

2.2.1 ESR

ESR

1 μ

(6)

g

(7)

e

2 , 2

(8)

α, β

2 Zeeman


19

2-3

ESR 2-3

CW-ESR

1 1

H

g

Zeeman Zeeman

2


20

R

A B

C, D

E, F


21

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


22

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


23

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


24

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]


25

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


26

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


27

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


28

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


29

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


30

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%


31

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


32

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


33

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


34

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%


35

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


36

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


37

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]


38

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.

Charlot M. F., Boussac A. and Blondin G., Towards a spin coupling model for the Mn4 cluster in

photosystem II, Biochimica et Biophysica Acta, 1708 (2005) 120-132. Cox N., Rapatskiy L., Su J. H., Pantazis D. A., Sugiura M., Kulik L., Dorlet P., Rutherford A. W., Neese

F., Boussac A., Lubitz W. and Messinger J., Effect of Ca2+/Sr2+ Substitution on the Electronic Structure of the Oxygen-Evolving Complex of Photosystem II: A Combined Multifrequency EPR, 55Mn-ENDOR, and DFT Study of the S2 State, Journal of the American Chemical Society, 133 (2011) 3635-3648.

Dismukes G. C. and Siderer Y., Intermediates of a Polynuclear Manganese Center Involved in

Photosynthetic Oxidation of Water, Proceedings of the National Academy of Sciences 78 (1981) 274-278.

Hara H., Kawamori A., Astashkin A. V. and Ono T., The distances from tyrosine D to redox-active

components on the donor side of Photosystem II determined by pulsed electron-electron double resonance, Biochimica et Biophysica Acta, 1276 (1996) 140-146.

Kessen S., Teutloff C., Kern J., Zouni A. and Bittl R., High-Field 2H-Mims-ENDOR Spectroscopy on

PSII Single Crystals: Hydrogen Bonding of YD●, Chemphyschem, 11 (2010) 1275-1282.

Kok B., Forbush B. and Mcgloin M., Cooperation of Charges in Photosynthetic O2 Evolution .1. A Linear

4step Mechanism, Photochemistry and Photobiology, 11 (1970) 457-&. Kulik L. V., Epel B., Lubitz W. and Messinger J., 55Mn pulse ENDOR at 34 GHz of the S0 and S2 states of

the oxygen-evolving complex in photosystem II, Journal of the American Chemical Society, 127 (2005) 2392-2393.

McEvoy J. P. and Brudvig G. W., Water-splitting chemistry of photosystem II, Chemical Reviews, 106

(2006) 4455-4483. Messinger J. and Renger G., Eds. Photosynthetic water-splitting Promary processes of

photosynthesis-part2: basic principles and apparatus. Cambridge, (2008). Royal Society of Chemistry.


39

Mino H., Astashkin A. V. and Kawamori A., An EPR and pulsed ENDOR study of the structure of tyrosine Z● in Tris-treated photosystem II, Spectrochimica Acta A, 53 (1997) 1465-1483.

Nagai H., Fukushima Y., Okajima K., Ikeuchi M. and Mino H., Formation of Interacting Spins on

Flavosemiquinone and Tyrosine Radical in Photoreaction of a Blue Light Sensor BLUF Protein TePixD, Biochemistry, 47 (2008) 12574-12582.

Nelson N. and Yocum C. F., Structure and function of photosystems I and II, Annual Review of Plant

Biology, 57 (2006) 521-565. Ono T. and Inoue Y., Effects of Removal and Reconstitution of the Extrinsic 33-Kda, 24-Kda and 16-Kda

Proteins on Flash Oxygen Yield in Photosystem-Ii Particles, Biochimica Et Biophysica Acta, 850 (1986) 380-389.

Pantazis D. A., Orio M., Petrenko T., Zein S., Lubitz W., Messinger J. and Neese F., Structure of the

oxygen-evolving complex of photosystem II: information on the S2 state through quantum chemical calculation of its magnetic properties, Physical Chemistry Chemical Physics, 11 (2009) 6788-6798.

Peloquin J. M., Campbell K. A., Randall D. W., Evanchik M. A., Pecoraro V. L., Armstrong W. H. and

Britt R. D., 55Mn ENDOR of the S2-state multiline EPR signal of photosystem II: Implications on the structure of the tetranuclear Mn cluster, Journal of the American Chemical Society, 122 (2000) 10926-10942.

Shen J. R. and Inoue Y., Binding and Functional-Properties of Two New Extrinsic Components,

Cytochrome-C-550 and a 12-kDa Protein, in Cyanobacterial Photosystem II, Biochemistry, 32 (1993) 1825-1832.

Shen J. R. and Kamiya N., Crystallization and the crystal properties of the oxygen-evolving photosystem

II from Synechococcus vulcanus, Biochemistry, 39 (2000) 14739-14744. Siegbahn P. E. M., A Structure-Consistent Mechanism for Dioxygen Formation in Photosystem II,

Chemistry-a European Journal, 14 (2008) 8290-8302. Siegbahn P. E. M., An Energetic Comparison of Different Models for the Oxygen Evolving Complex of

Photosystem II, Journal of the American Chemical Society, 131 (2009) 18238-18239. Siegbahn P. E. M., Structures and Energetics for O2 Formation in Photosystem II, Accounts of Chemical

Research, 42 (2009) 1871-1880. Stich T. A., Yeagle G. J., Service R. J., Debus R. J. and Britt R. D., Ligation of D1-His332 and

D1-Asp170 to the Manganese Cluster of Photosystem II from Synechocystis Assessed by Multifrequency Pulse EPR Spectroscopy, Biochemistry, 50 (2011) 7390-7404.

Styring S. and Rutherford A. W., In the Oxygen-Evolving Complex of Photosystem-Ii the S0 State Is

Oxidized to the S1 State by D+ (Signal-Iislow), Biochemistry, 26 (1987) 2401-2405. Umena Y., Kawakami K., Shen J. R. and Kamiya N., Crystal structure of oxygen-evolving photosystem II

at a resolution of 1.9 Å, Nature, 473 (2011) 55-U65. Yeagle G. J., Gilchrist M. L., McCarrick R. M. and Britt R. D., Multifrequency pulsed electron

paramagnetic resonance study of the S2 state of the photosystem II manganese cluster, Inorganic Chemistry, 47 (2008) 1803-1814.


40

7

Faisal Hammad Mekky Koua

1

1 4

G


2012 º Ø 1= emercury.yukawa.kyoto-u.ac.jp/~bussei.kenkyu/pdf/02/2/...2.1.2 4ß(ò$Î#Õ q ö b 6...

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