光合成におけるエネルギー生産について
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1
2 CO 2
3 CO 2
4 RuBisCO
5 phytochromobilin Pr Pfr
6 CO 2
7
8 1 CO2
9 a b
10 b CHO a a -
11 H+ H+ e - e - Mn Yz e - Q B P680 Q A Mn Yz P680 Q A Q B e ē - e - Mn Yz P680 Q A Q B Mn e - Yz P680 e - Q A Q B H 2 O Mn Yz e - P680 e - Q A Q B O 2 H + Mn Yz e - P680 e - Q A Q B PSI
12 Chl Chl Light-harvesting chlorophyll, LHC Chl Chl Chl Chl Chl Chl Chl NADPH Chl Chl Chl H + (CH 2 O) n CO 2 Chl Chl Chl Chl Chl a Chl H + b Chl Chl Chl ATP Chl Chl Chl H + γ β δ α β α c β ε α ADP + Pi ATP ATP
13 - 800 mv Q A /Q B PS II mvh 2 O ½O 2 Z Scheme Fe-S b6/f mv NADP NADP NADPH mv PS I
14 CO 2
15 CO 2 /O 2 RuBP GAP SBP F6P R5P R5P S7P Xu5P SBP Ru5P E4P FBP FBP GAP Xu5P DPGA Ru5P RuBP RuBisCO PGA TP DAHP TP CO 2 O 2 PGA glycolate GAP RuBP glyoxylate glycine P-glycolate PCO serine glycerate CO2O2 hydroxypyruvate O 2 ATP, NADPH
16 RuBisCO CO2 O2 CH2OPO3 - C=O HC-OH HC-OH CH2OPO3 - RuBP C-OH C-OH HC-OH * CH2OPO3 - CH2OPO3 - O 2 CO2 HO - HO - CH2OPO3 - CH2OPO3 - C-COO * - HO C-COO * - C=O - HO-C-OH HC-OH CH2OPO3 - CH2OPO3 - C-OO - C=O HC-OH CH2OPO3 - HC-OH CH2OPO3 - CH2OPO3 - HO C-OO - - HO-C-OH HC-OH CH2OPO3 - CH 2OPO3 - HC-OH * COO - + COO - HC-OH CH2OPO3 - PGA CH2OPO3 - COO - + COO - HC-OH CH2OPO3 - PGA
17 Ci Sr = Vcmax/Kcm)/(Vomax/Kom) CO 2 O 2 CO2 C C SS CO2 O2 Ci Ci Sr
18 -1,6- C6-6- C6 C3) C3) C12) C12) Cn
19 RuBisCO GAP R5P R5P S7P Xu5P SBP SBP Ru5P Xu5P Ru5P RuBP F6P E4P FBP FBP GAP CO 2 O 2 DPGA RuBP RuBisCO PGA TP DAHP TP PGA glycolate GAP glyoxylate glycine P-glycolate PCO serine glycerate O 2 hydroxypyruvate
20 CO2 photons
21 CO 2
22 2000 CO2
23 CO 2 /O 2 CO 2 SuperRuBisCO CO 2 (ppm)
24 PNAS (1997) Science (2003) (rbcs) Cross-talk mrna (rbcl) RuBP mrna worse RuBisCO Form I of cyanobacteria, green algae and higher plants Form II k cat :3-8 s -1 site -1 S rel :10-90 BBRC (2003) JMB (2002) JBC (1997) PCR/PCO cycles CO2/O2 RuBP better RuBisCO Form I of Galdieria and some red algae S rel :240 Engineered Form I k cat :16 s -1 site -1 FEBS Lett. (2002) JB (2001) JBC (1999) BBRC (1997)
25 CO 2 O 2RuBisCO O2 T65 K177 E60 N123 R295 CO 2 RuBisCO RuBisCO CO 2 O 2 RuBisCO RuBisCO loop 6 CO 2 O 2 O 2
26 1 CO CO 2
27 CO 2 CO 2 CO 2
28 Late Embryogenesis Abundant (LEA)
29 P680 II ATP 2 H + NADPH NADP H + CF1 α β LHC2 Phe QA QB P680 D2 D1 Mn 33kD 1/2 O2 27kD H2O + 17kD 2 H + PCR/PCO PQ LHC2 PQ PQ 2 H + Cyt.b6 Cyt.f RuBisCO Fd FNR A Fe-S CF0 P700 PC 3 H + 3 H + ADP + Pi P700 I
30 OH O 2 - SOD OH NADP GSH H 2 O 2 V.C GSSG NADPH GR GPX APXMDARDHAR GR NADPH GSSG H 2 O MDV.C GSH NADP V.C
31 A B 1 2 Nos-pro LB NPTII V S C M S Q GTC AGT TGC ATG TCG CAA transit peptide HindIII Nos-ter pro KatE tomato rbcs3c gene StuI/SstI Nos-ter RB 93kD Wild tobacco Transformants
32 µmol/mg protein/min µmol/mg chlorophyll/h nmol/mg protein/min cyt chl wild transformant CO2 fix Ru5PK APX 0 hr 48 hr 0 hr 48 hr NADP + O2 e- Photosystems 1/2 O2 PCR/PCO Cycles GSH/ GSSG O2 - /H2O2 H2O DHAR? H2O Ascorbate APX Monodehydro -ascorbate Dehydroascorbate H2O Chloroplast
33
34
35 1 Craterostigma
36
37
38
39 H 2 N : NH 2 C=O NH CH 2 CH 2 CH 2 CH COOH #. #. # % Citrulline Arginine Glutamic acid Glutamine Alanine Serine Aspartic acid (M -1 s -1 ) ( ) X 10 9 ( ) X 10 9 ( ) X 10 8 ( ) X 10 7 Kawasaki et. al., (2000) Plant Cell Physiol., 41, Akashi, et al., (2001) FEBS Lett. 508,
40 b 561 ε-
41 CO 2
42 rbcl mrna RNA/ 3500 DNA DNA Gppp AAAAA DNA10,000 copies/cell Gene Dosage
43 mrna CO2
44
45 CO 2
46
47
48
49 CO 2 O 2 R5P R5P S7P Xu5P SBP SBP Ru5P Xu5P Ru5P RuBP glyoxylate GAP E4P F6P FBP FBP DPGA CO 2 O 2 RuBP RuBisCO PGA PGA GAP GAP glycolate glycine P-glycolate PCO serine glycerate hydroxypyruvate TP DAHP TP
50 UP RuBisCO CO2 UP
51 RuBisCO
p.1127 Chapter 22 Amino Acids, Peptides, and Proteins p.1128 p.1128 Amino acid Peptide Amino acid p.1132 p.1128 p.1129 p.1129 p.1129 p.1130 p.1130 p.1130 p.1130 p.1130 p.1130 p.1130 p.1130 p.1130 p.1130
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