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Objectives
• Discuss photosynthesis at the molecular level in
terms of the structural changes, enzymes, co-
factors, cell localization and regulation.
• Discuss the energetics involved in the processes
of photosynthesis.
• Compare and contrast photosynthesis in C3, C4
and CAM plants

CHEM 160
PHOTOSYNTHESIS 1 2

Photosynthesis Photoautotrophs
• The process by which autotrophic organisms
convert CO2 into sugars with the use of energy
coming from sunlight
• Basically involves conversion of light energy
into chemical energy algae
Plants

Unicellular protists Cyanobacteria

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Chloroplasts Chloroplasts
• Site of photosynthesis

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Chloroplasts Chlorophylls
• Excellent light harvesters due to the presence
of conjugated double bonds

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Other light harvesting pigment Other light harvesting pigment

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Absorption of visible light by Why are Plants Green?

photopigments

Different pigments absorb light differently

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Two Phases of Photosynthesis Overview of Photosynthesis

1. Light Reactions
 Occurs in the thylakoid
 Light energy is converted to chemical
energy
 ATP and NADPH are produced
2. Dark Reactions
 Occurs in the stroma
 CO2 is fixed and reduced to form sugars
 Uses ATP and NADPH
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Photosystems (PS)
 Multi-subunit protein complexes embedded in the
thylakoid membrane responsible for light-driven
photosynthesis

 Two types of photosystems cooperate in the light

LIGHT REACTIONS OF reactions (PS I and II)

PHOTOSYNTHESIS
PS II PS I

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Photosystems Light Reactions – Z Scheme

X-ray structure of PSII from T. elongatus.

Cyt B
complex
Pq

Pc

Cyanobacteria photosystem II 18 19

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Photosystems Photosystems
Each photosystem has Photochemical
reaction center
 reaction center
complex or
photochemical are specialized to
reaction center transduce light into
chemical energy.
 Light-harvesting
complexes or
antenna molecules

Campbell, 2009 Campbell, 2009

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Photosystems Photosystems
Photochemical Light-harvesting
reaction center complexes or
antenna molecules.
 Special pair of
chlorophyll a absorb light energy and
molecules transmit it rapidly and
efficiently to the
reaction center

 Primary electron
Campbell, 2009 acceptor Campbell, 2009

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Excitation of e− in Chlorophyll

Campbell, 2009

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4

Light Reactions
• Linear; NADPH and ATP are both produced
Cyt B
complex
Pq
Pc
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Light Reactions - Noncyclic e- flow
It excites P680 e- which is
captured by the primary e-
Cyt B
acceptor leaving an e- hole
complex in the P680
Pq
Pc
Electron will then be
What will replace an transferred to PSI via an
e- hole in the P680? electron transport chain
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Light Reactions - Noncyclic e- flow
1. Excitation of e− in PS II
It excites P680 e- which is
captured by the primary e-
Cyt B
acceptor leaving an e- hole
complex in the P680
Pq
Pc
Electron will then be
What will replace an transferred to PSI via an
e- hole in the P680? electron transport chain
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Light Reactions - Noncyclic e- flow
Excitation of e− in PS II
A photon of light is absorbed
by a chlorophyll in a light
harvesting complex converting
Cyt B it to a higher E state .
complex
Pq
Pc
The excitation E will then be
transferred to neighboring
chlorophyll until it reaches one
of the two P680 chlorophyll a
molecules
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Electron transfers in PS I
Plastocyanin transfers
electrons to P700
Cyt B
Pq complex
Pc
Excited electrons in
P700 is transferred to
ferredoxin then to
NADP+ reductase
NADP+ reductase uses
high energy electrons
to reduce NADP+ to
NADPH 29
What will replace an e- hole in the P680?
An enzyme splits a water
molecule into two electrons,
+
Cyt B two H , and an oxygen atom.
Pq complex
The electrons are supplied
onePcby one to the P680, each
replacing an electron lost to
the primary electron acceptor.
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Electron Transport Chain Comparison of chemiosmosis in

mitochondria and chloroplasts
ETC components:

• Plastoquinone
Cyt B
Pq complex • Cytochrome
complex
Pc
• Plastocyanin

ETC generates a proton

gradient that drives
ATP synthesis

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Light Rxns and ATP Synthesis Cyclic Electron Flow

• Occurs when NADP+ is not available
• e− from Fd is transferred back to ETC between PS II and PS I
• Only ATP is generated.

Campbell, 2009

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Dark Reactions
• Occurs in the stroma of the chloroplast
• Light is still needed for the activation of some
involved enzymes.
DARK REACTIONS OF CALVIN-BENSON CYCLE or
C3 PHOTOSYNTHESIS
PHOTOSYNTHESIS • Reduction of carbon from CO2 in a more
reduced state as hexose
• Uses NADPH and ATP produced from the light
reactions

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Calvin Cycle 1. CO2 Fixation

• Fixation of CO2 with ribulose-1,5-bisphosphate
(RubP) to form 2 molecules of 3-
phosphoglycerate (3-PGA)
• Catalyzed by RuBisCO

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1. CO2 Fixation 2. Reduction of 3-PGA to GAP

ATP ADP
RuBisCO O O PO 32-
O O
C C
• Ribulose-1,5-bisphosphate carboxylase-oxygenase H OH H OH
phosphoglycerate
• Most abundant enzyme on earth CH 2O PO 32- kinase CH 2O PO32-
3-phosphoglycerate 1,3-bisphosphoglycerate
• Requires Mg2+ as co-factor
NADPH
glyceraldehyde-3-P dH

NADP+ + Pi

O H
C
H OH
CH 2O PO 32-
glyceraldehyde-3-P

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3. Regeneration of RubP Stoichiometry

• Proceeds in the
same fashion as
the reverse of the
non-oxidative
stage of PPP

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Stoichiometry RuBisCO: Oxygenase Reaction

• To fix 1 molecule of CO2:
• 3 ATP and 2 NADPH molecules are required
• To synthesize a glucose molecule:
• 6 rounds of Calvin cycle
• 18 ATP and 12 NADPH molecules are
required
• Rubisco also catalyzes a wasteful oxygenase
reaction when [CO2]/[O2] is low
• Happens under hot dry condition. Stomata
cannot open to allow entry of CO2
• A phenomenon called PHOTORESPIRATION

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RuBisCO: Oxygenase Reaction RuBisCO Reactions

www.tutorvista.com www.tutorvista.com

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RuBisCO: Oxygenase Reaction C4 Photosynthesis

• Why wasteful? • Alternative strategy of some tropical plants
• Consumes ATP and NADPH in the process (eg. corn, sugarcane) that are usually exposed
itself and in the detoxification of the side to hot dry environment
product NH3. • Prevents oxygenase activity of RuBisCO by
• CO2 that has already been fixed is released maintaining a high [CO2]
again as CO2 • Uses a carbon-fixing enzyme PEP carboxylase
• O2 is consumed. in tandem with RuBisCO
• Uses a unique anatomical difference to ensure
optimal C-fixation

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Anatomy of C3 and C4 Leaf C4 Photosynthesis

1. CO2 is added to PEP
C3 C4
to form OAA (PEP
carboxylase)
2. MC export OAA to the
BSC through malate
3. In the BSC, malate is
broken down to CO2
and pyruvate.
• Close association of bundle
• Scattered mesophyll cells
sheath cells (BSC) with 4. CO2 is converted to
(MC) sugar via normal
mesophyll cells
• Only MC contain
• Both MC and BSC contain
Calvin Cycle
chloroplast
chloroplast 50 51

Advantages of C4 over C3 CAM Plants

Plants • CAM - Crassulacean Acid Metabolism
1. No photorespiration • Observed in Crassulaceae (succulents)
2. At ⬇[CO2], PEP carboxylase concentrates CO2 in • Thick cuticles
BSC to favor carboxylase reaction of RuBisCO • High water content
3. Close association of BSC and MSC allow more • Lower number of stomates
efficient transfer of intermediates
4. Both MC and BSC contain chloroplast

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CAM Plants
• During DAYTIME
• Stomates are closed to prevent evaporation
of H2O; no CO2 can enter the plant
• During NIGHTIME
• Stomates are open; CO2 is fixed via C4
pathway
• The following DAYTIME
• Malate is decarboxylated and CO2 is fixed
via the Calvin Cycle.

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