Dark Reactions

Cyclic photosphorylation
generates
Stromal side
ATP
Fd

PQ

PQ

P700
PSII PSI

Thylakoid space side

Non-Cyclic photosphorylation
generates ATP and NADPH
Stromal side NADP+ NADPH

Q
Fd
PQ

PQ

P680 P700
PSII PSI

H20 2H+ & 2e- & 1/2 O2 Thylakoid space side

Light-independent reactions
(Dark Reactions)
The Calvin/Benson/C3 cycle

• Has 3 phases:
1. carbon fixation phase
2. reduction of CO2 phase
3. regeneration of RuBP phase
carbon fixation phase
• each CO2 molecule is attached to a five-
carbon sugar, ribulose bisphosphate
(RuBP)
• catalyzed by RuBP carboxylase or rubisco
• six-carbon intermediate splits in half to
form two molecules of 3-phosphoglycerate
reduction of CO2
phase
• each 3-phosphoglycerate receives another
phosphate group from ATP to form 1,3
bisphosphoglycerate.
• pair of electrons from NADPH
reduces,Gain, each 1,3
bisphosphoglycerate to G3P.
– Electrons from NADPH change a
carboxyl group to a carbonyl group.
regeneration of RuBP
phase
• regeneration of the CO2 acceptor (RuBP),
these five G3P molecules are rearranged
to form 3 RuBP molecules.
• cycle must spend 3 more molecules of
ATP (one per RuBP) to complete the cycle
and prepare for the next.
Hatch & Slack/C4 cycle

• Inefficient as RUBISCO can act as both a

carboxylase and as an oxygenase.
• Oxygenase activity leads to loss of carbon that
has already been fixed.
• Some plants have an alternative pathway where
CO2 is first fixed into C4 organic acids (C4
pathway) and then liberated later to undergo the
Calvin/Benson cycle (C3 metabolism).
• 2 types of C4 cycle:
1. C3 and C4 metabolism are separated in space
(different cells) e.g. sugar cane
Characteristic aggregation of cells around the
vascular bundles - Krantz morphology (
C4 cycle)
2. C3 and C4 metabolism are separated in time
(same cell=mesophyl) e.g. pineapple (
CAM cycle).
Comparison of C3 & C4 leaves
• C3 - note the lack of • C4 - note the extensive
chloroplasts in the chloroplasts in the
bundle sheath bundle sheath
 CO2
Separation
in space (C4 cycle)
PEP (3C) Mesophyll cell
OAA (4C)
ADP surrounding
ATP bundle sheath

Malat Pyruvate (C3)

CO2
C3 Bundle sheath cell
sugar

Vascular tissue
 CO2
Separation
in time (CAM cycle)
PEP (3C) In the dark
OAA (4C)
ADP
ATP

Malat Pyruvate (C3)

CO2
C3 In the light
sugar

Mesophyl
Comparison of C3 & C4 leaves
Details of C4 cycle
Details of CAM cycle