Dark Reaction Topics

CO2 Fixation

Calvin Cycle

Starch and Sucrose Biosynthesis

C3 vs C4 Plants
 Dark Reaction
Glucose
Glucose Oxidation
Oxidation Exergonic
Exergonic

CC66H
H1212O
O66 ++ 6O 6CO
6CO22 ++ 6H O G
6H22O Go’==-2,860
-2,860kJ/mol
o’
6O22 kJ/mol

Dark
Dark Reaction
Reaction Endergonic
Endergonic

6CO
6CO22 ++ 6H
6H22O CC66H
H1212O 6O22 G
O66 ++ 6O Go’==++2,860
2,860kJ/mol
o’
O kJ/mol
 CO2 Fixation (One Enzyme)
CH2OP Enediol CH2OP CO CH2OP
C=O  electrons C-OH
2

HO-C-CO2
C-OH C-OH C=O
C-OH C-OH C-OH
CH2OP CH2OP CH2OP
Ribulose 1,5bisPO4
H 2O
CH2OP CH2OP CH2OP
HO- C-CO2 H +
HO- C-CO2 HO- C-CO2
H ..
2, 3PGA + HO-C-OH
COO- HO-C-OH H C-OH
H C-OH H C-OH CH2OP
CH2OP CH2OP
Ribulose 1,5 bisPO4 carboxylase or RUBISCO
 The Calvin Cycle in
Photosynthesis
Rule: The Calvin cycle is the mechanism for turning CO2
into glucose
Rule: The cycle fixes 6 CO2 to 6 ribulose 1,5-bisPO4
acceptor molecules, makes glucose with 6 of the
carbons, and returning the rest to continue the cycle.

Rule: The driving force is to synthesize the 5 carbon

product ribulose 1,5-bisPO4
 Calvin Cycle Strategy
(Ideal)
6CO2 Glucose
(Real)
6CO2 + 6RuBP G6P + 6RuBP
6C 30C 6C 30C
(Mechanism)
6CO2 + 6RuBP 12, 3PGA
6C
12, 3PGA G6P + 2F6P + 4G3P + 2DHAP
2E4P + 2SHBP 4Xu5P + 2Ru5P
Intermediates 20C 10C
Final
 Carbon Balance in
12 3-PGA 36c
36c
ATP Calvin Cycle
ADP

12 1,3 bisPO4 glycerate 36c

36c
NADPH + H+
NADP+
24c
24c 6c
6c
12 glyceraldehyde-3-PO4 DHAP Calvin
Calvin

Calvin
Calvin 3 Fructose 1,6bisPO4 18c
18c
12c
12c
2 Fructose-6-PO4 Calvin
Calvin
12c
12c
Glucose-6-PO4
6c
6c
 C TK and TA Reactions
C=O
C TK C
C=O
C CHO HO C Ru5P
C C C
CH2OP CH2OP CH2OP
F6P G3P Xu5P
CH2OP Pi
C=O TK C CHO
HOC C=O C
CHO TA COH HO C + C
CH2OP C C
C C CHO
C=O C CH2OP CH2OP
C C Xu5P
CH2OH CH2OP CH2OP CH2OP R5P
DHAP E4P S1,7BP G3P Ru5P
 Finishing Touches with 5 Carbon
Rule: In the Calvin cycle 5C sugars are never donor or
acceptor substrates for transketolases or transaldolases

CH2OH CH2OH CH2OP

C=O Epimerase C=O ATP C=O
HO-C C-OH C-OH
C-OH C-OH C-OH
CH2OP Kinase
CH2OP CH2OP
Xu5P Ru5P
CHO CH2OH RuBP
C-OH Isomerase C=O
C-OH C-OH ATP 30 carbons are
returned to
C-OH C-OH
CO2 fixation
CH2OP CH2OP
R5P Ru5P
 G3P
3C DHAP
G3P
3C
3C 3C G3P
DHAP 3C
6C 4C 7C
F6P E4P SBP
5C
5C
Xu5P 5C
R5P
Xu5P
 Synthesis of Sucrose and Starch
F6P  G6P  G1P Cytosol
ATP UTP
PPi PPi
ADP-glucose UDP-glucose
(glucose)n fructose-6-PO4

ADP + (glucose)n+1 UDP + Sucrose-6-PO4

Stroma Starch
H2O
(amylose) Pi
Sucrose
Thank goodness for
photosynthesis
 Control of the Calvin Cycle
ΔGo’ ΔG
Phosphoribokinase -21.8 -15.9
Ribulose bisPO4 carboxylase -35.1 -41.0
Phosphoglycerate kinase + +18.0 -6.7
glyceraldehyde-3-PO4 dehydrogenase
Triose phosphate isomerase -7.5 -0.8
Aldolase -21.8 -1.7
Fructose bisphosphatase -14.2 -27.2
Transketolase +6.3 -3.8
Aldolase -23.4 -0.8
Sedoheptulose bisphosphatase -14.2 -29.7
Transketolase +0.4 -5.9
Phosphopentose epimerase +0.8 -0.4
Ribose phosphate isomerase +2.1 -0.4

Each
Eachcontrolling
controllingenzyme
enzymeisisitself
itselfregulated
regulatedby
bylight
light
either
eitherthrough
throughpH
pHor orMg
+2
Mg+2
 Control of Flux in Plants

Glyceraldehyde-3-PO4

Pi
Glyceraldehyde-3-PO4 NADP+
NADPH
dehydrogenase
1,3-Bisphosphoglycerate

Phosphoglycerate ADP Does the Calvin

kinase ATP cycle occur in the
dark?
3-PGA
NO
Calvin
Cycle
 CH2OPO3
Therefore
ThereforeCO CO22 Binds
Bindstightly
tightlytoto
HO-C-CO2 RUBISCO
fixation
fixationcannot
cannot RUBISCO
H-C-OH when
occur
occurin
inthe
the whenthere
thereisis
dark H-C-OH no
nolight
light
dark
CH2OPO3

2-Carboxyarabitol-1-PO4
Active*

PS1*RED FdOX FdRRED Thioredoxin 1,7SH

BisPtase
SH
SH SH S S

PS1OX FdRED FdROX Thioredoxin 1,7 BisPtase

SH SH
S S
Inactive
Sedoheptulose
Sedoheptulose1,7-bisphosphatase
1,7-bisphosphatase
isisactivated
activatedby
bylight
light Text
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