3/13/2018

Photosynthesis

Breakdown of Photosynthesis 1.Light Reaction (Electron Flow)

Two main parts (reactions):
1. Light Reaction or Light Dependent Reaction
Produces energy from solar power (photons)
in the form of ATP and NADPH
2. Calvin Cycle or Light Independent Reaction
or Carbon Fixation or C3 Fixation
Uses energy (ATP and NADPH) from light
rxn to make sugar (glucose)
Occurs in the Thylakoid membranes
During the light reaction, there are two possible
routes for electron flow
A.Cyclic Electron Flow
B.Noncyclic Electron Flow

A. Cyclic Electron Flow Cyclic Electron Flow

Occurs in the thylakoid membrane
Uses Photosystem I only Primary
Electron e-
P700 reaction center- chlorophyll a Acceptor

Uses Electron Transport Chain (ETC)

e- e- ATP
Generates ATP only produced
Photons
by ETC
ADP + P ATP P700
e-
Accessory
Pigments

Photosystem I

1
 3/13/2018

B. Noncyclic Electron Flow Noncyclic Electron Flow

Occurs in the thylakoid membrane Primary

Uses PS II and PS I
Electron
Acceptor
2e- Enzyme
Reaction
P680 rxn center (PSII) - chlorophyll a Primary
Electron 2e-
Acceptor ETC 2e-
P700 rxn center (PS I) - chlorophyll a
Uses Electron Transport Chain (ETC) 2e- NADPH
Generates O2, ATP and NADPH 2e- P700
Photon ATP
P680 Photon
H2O
Photosystem I
1/2O2 + 2H+ Photosystem II

PS II
Noncyclic Electron Flow Absorption of light by PS II => provide energy, allow
photooxidation of H2O (final e source)
ADP + P  ATP ChlII + hv (680nm)  ChlII*
(Reduced)
Where,
NADP+ + H  NADPH ChlII == P680 Chl
(Reduced) ChlII*
strong reducing agent, pass e to ChlI+ through an
Oxygen comes from the splitting of H2O, not CO2
e transport chain coupled to the phosphorylation
of ADP  ChlII+
H2O  ½ O2 + 2H+ Energy for rxns to allow e flow from PS II to
PS I – supplied by absorption of light by ChlII
(Oxidized) ChlII+ - lack 1e, become strong oxidizing agent
to oxidize H2O to O2

Next e acceptor is plastoquinone (PQ)

PS II: H2O Is Split To Produce O2
Oxidation of H2O to produce O2 – ultimate e
source in PS
e passed from PS II to PS I by e transport chain
Oxidized ChlI+ produced by PS I – reduced by e
donated by PS II
ChlII* - reducing agent, passes e to a primary
acceptor, Pheophytin (Phe)
e transport chain of the thylakoid membrane
linked 2 PS consist:
Phe, PQ, complex of plant cytochromes (b6 - f
complex), plastocyanin (copper-containing
protein), & oxidized P700 (ChlI+)
ATP generated in this e transport chain
When oxidized P700 (or PSI) (ChlI+) accept e,
become reduced (ChlI)
And rxn center Chl of PS II (P680), oxidized
state, gains e, become reduced as result of
oxidation of H2O
2 H2O + 4 ChlII+  O2 + 4 H+ + 4 ChlII

2
 3/13/2018

PS I
Absorption of light by Chl => supply energy, allow
photoreduction of NADP+
4 e required for H2O oxidation ChlI + hv (700nm)  ChlI*
ChlII+ (P680+) can accept only 1e a time Where,
Net rxn for 2 PS == flow of e from H2O to NADP+ hv = light absorbed
ChlI & ChlI* == P700 Chl in ground (unexcited) and excited states,
2 H2O + 2 NADP+  O2 + 2 NADPH + 2 H+ respectively

ChlI*
strong reducing agent, easily give e to another
substance
give 1e to NADP+, produce NADPH
NADP+ + e  NADPH
lack 1e become ChlI+
the needed e supplied by PS II

PS I: Reduction of NADP+ ChlI* + ferrodoxin (oxidized) ChlI+ +

ChlI* give 1e to be transferred
e transfer mediated by processes take place in
rxn center
Next e acceptor – bound ferrodoxin, an iron-
sulfur protein
Bound ferrodoxin passes e to a molecule of
soluble ferrodoxin
Soluble ferrodoxin reduced by FAD enzyme
called ferrodoxin-NADP reductase
Reductase reduces NADP+ to NADPH
ferrodoxin (reduced)
2 Ferrodoxin (reduced) + H+ + NADP+
Ferredoxin-NADP 2 ferrodoxin (oxidized) + NADPH
reductase
ChlI* donates 1e to ferrodoxin
1e from 2 ferrodoxin is required for production
of NADPH
Oxidized Chl (ChlI+) => lost 1e, colorless
the e needed to reduce ChlI+ supplied by PS II

Photophosphorylation & Proton

Gradient In Chloroplasts

3
 3/13/2018

Photosynthetic e transport pumps H+ out of the

stroma to the intrathylakoid space to form the
proton gradient
High pH in the stroma, low pH in the
intrathylakoid space
The flow of H+ back to the stroma through the
ATP synthase provides the energy for synthesis
of ATP from ADP and Pi

Dark Rxn: CO2 Fixation

CO2 fixation take place in stroma
Overall rxn
ATP
6CO2 + 12 NADPH C H O + 12 NADP+
Enzymes 6 12 6
Net rxn: 6 molecules of CO2 produce 1 molecule
of glucose

Reaction Processes Calvin Cycle

Carboxylation of 6 molecules of a 5-C key
intermediate, ribulose 1,5-bisphosphate
Formation of 6 molecules of unstable 6-C
intermediate
Splitting to 12 molecules of 3-phosphoglycerate
Production of glucose from 2 molecules of 3-
phosphoglycerate
Generation of 6 molecules of ribulose 1,5-
bisphosphate from the remaining 10 molecules of
3-phosphoglycerate
overall rxn pathway is cyclic
Calvin cycle

4
 3/13/2018

O O
CH 2 O P O CO2
CH 2 O P O
Calvin Cycle C O O COO
-
C OH O
H C O C O
First step H C OH O H C O O
- -
CH2 O P O
1. Condensation of ribulose 1,5-bisphosphate with CH2 O P
-
O
O
-

CO2 O

Ribulose 1,5-bisphosphate 2-carboxy-3-ketoribitol 1,5-bisphosphate

2. Formation of a 6-C intermediate, 2-carboxy-3-
ketoribitol 1,5-bisphosphate H2O

3. Hydrolyzation to give 2 molecules of 3- Rxn of ribulose 1,5-

phosphoglycerate
bisphosphate with CO2
catalyzed by enzyme ribulose 1,5- O
bisphosphate carboxylase produces 2 molecules
-
C O
located on the stromal side of the thylakoid of 3-phosphoglycerate
H C OH O
membrane CH2 O P O
-

mw  560,000; consists of 8 large subunits & O

-

8 small subunits TWO 3-phosphoglycerate

O
ATP O 2-
C O- PO3

H C OH O C O

CH2 O P O- H C OH O
-
Second step O- ADP CH2 O P O
O-
1. Reduction of 3-phosphoglycerate to form 3-phosphoglycerate 1,3-bisphosphoglycerate

glyceraldehydes 3-phosphate NADPH

2. Require NADPH NADP +

+ Pi
3. NADPH & ATP generated in light rxn provide Reduction of
energy for the dark rxn 3-phosphoglycerate to O
glyceraldehyde 3-phosphate C H
H C OH O
-
CH2 O P O
O-
Glyceraldehyde 3-phosphate

Glucose
Production of
glucose from
Glucose 6-phosphate
glyceraldehyde
Third step (2 alternative ways) 3-phosphate
A. Production of 6-C sugars Fructose 6-phosphate

1. Conversion of G3P to dihydroxyacetone

Fructose 1,6-bisphosphate
phosphate (DP)
2. DP reacts with G3P to form glucose 6- Glyceraldehyde 3-phosphate Dihydroxyacetone phosphate

phosphate (G6P) 2 NADP+ + Pi

3. Formation of glucose from G6P 2 NADPH

The use of NADPH
2 1,3-bisphosphoglycerate & ATP generated in
light rxn to provide
2 ADP

2 ATP
energy for dark rxn
2 3-phosphoglycerate

5
 3/13/2018

6 CO2
CO2 Fixation
6 C-C-C-C-C-C
B. Regeneration of ribulose 1,5-bisphosphate RuBP
The enzyme that catalyzes
Carboxylase
Process involve 4 steps: (rubisco) this reaction is ribulose
1. Preparation biphosphate carboxylase
6 C-C-C-C-C (rubisco)
2. Reshuffling
3. Isomerization CO2 fixation refers to bonding CO2 to an
4. Phosphorylation organic molecule to make a larger molecule.
Each CO2 is bonded to ribulose biphosphate
(RuBP).
C5 + CO2  C6

C3 Photosynthesis – Calvin Cycle Calvin Cycle

6 CO2
6 CO2
6 C-C-C-C-C-C 6 C-C-C-C-C-C
RuBP RuBP
Carboxylase Carboxylase
(rubisco) (rubisco)

RuBP 6 C-C-C-C-C RuBP 6 C-C-C-C-C PGA 12 C-C-C

PGA 12 C-C-C

PGAL
12 C-C-C
Each of these 6-carbon
compounds splits to form two 3- The two molecules of PGA
carbon compounds called are reduced to form PGAL
phosphoglycerate (phosphoglyceraldehyde)

Calvin Cycle Calvin Cycle

6 CO2 6 CO2
6 C-C-C-C-C-C 6 C-C-C-C-C-C
RuBP RuBP
Carboxylase Carboxylase
(rubisco) (rubisco)

RuBP 6 C-C-C-C-C RuBP 6 C-C-C-C-C PGA 12 C-C-C

PGA 12 C-C-C
6 ADP + P
6 ATP
Two of the PGAL are used to form glucose
12 ATP 10 C-C-C phosphate, then glucose 12 ATP
PGAL PGAL
12 C-C-C 12 ADP + P 12 C-C-C 12 ADP + P

12 NADPH 12 NADPH

12 NADP+ C-C-C-C-C-C Glucose 12 NADP+

6
 3/13/2018

Calvin Cycle Calvin Cycle

6 CO2 6 CO2

6 C-C-C-C-C-C 6 C-C-C-C-C-C
RuBP RuBP
Carboxylase Carboxylase
(rubisco) (rubisco)

RuBP 6 C-C-C-C-C RuBP 6 C-C-C-C-C

The remaining PGA 12 C-C-C PGA 12 C-C-C
6 ADP + P 6 ADP + P
10 PGAL are This process requires energy in the
6 ATP rearranged to 6 ATP form of ATP.
12 ATP
form 6 RuBP
10 C-C-C 10 C-C-C 12 ATP
PGAL 12 ADP + P PGAL
12 C-C-C 12 C-C-C 12 ADP + P
12 NADPH
12 NADPH
C-C-C-C-C-C Glucose 12 NADP+
C-C-C-C-C-C Glucose 12 NADP+

Overview of Photosynthesis