1

7.0: Conversion of Light Energy to

Chemical Energy – Light there be life!

Key Concepts (the big picture):

• Photosynthesis takes place in the chloroplasts

(and many bacteria).
•Photoreceptive compounds (eg. Chlorophyll(s))
within the chloroplast are grouped together with
other compounds, to form photosystems, and it
is these complexes which absorb light.

1
 Key Concepts (the big picture) cont:

The processes within photosystems are called the

light reactions. These reactions include:
- splitting of water to produce O2 and H+
- NADP+ reduction to produce NADPH
- H+ gradient generation to power ATP synthesis
(yes chloroplasts undertake oxidative
phosphorylation!)

WHAT YOU NEED TO KNOW!! You should:

• describe the two photosynthetic stages and the

overall reaction of photosynthesis
• explain the structure of the chloroplast and the
molecules involved in light absorption
• understand the reactions taking place during the
light reactions
• discuss how the absorption of light is coupled to ATP
synthesis

2
5

Photosynthetic light harvesting can achieve a

quantum efficiency that approaches 100% (that is,
the conversion of 100 photons of light into 100
chemically available electrons), and yet it displays
notable robustness in the face of ever-changing
external light conditions.

But a bigger question is why are plants green!

3
 https://www.quantamagazine.org/why-are-plants-
green-to-reduce-the-noise-in-photosynthesis-
20200730/

4
9

Mathematical model predicted that the absorption peaks of

chlorophyll a and b, which green plants use to harvest red
and blue light. It appears that the photosynthesis
machinery evolved not for maximum efficiency but
rather for an optimally smooth and reliable output.

10

5
 A tentative model for the organization of the thylakoid membrane
This is a good summary so learn this

STROMA
(low H+ concentration) Cytochrome
Photosystem II Photosystem I
complex
4 H+ Light NADP+
Light reductase
Fd 3
NADP+ + H+

Pq NADPH

e– Pc
e– 2
H2O
THYLAKOID SPACE 1 1/
2 O2
(high H+ concentration) +2 H+ 4 H+

To
Calvin
Cycle

Thylakoid
membrane ATP
STROMA synthase
ADP
(low H+ concentration)
+ ATP
Pi
H+

11

The site of photosynthesis in a plant

(bacteria and algae do this as well)

The thylakoid membrane is impermeable to most ions and

molecules (not Mg2+ and Cl- however)
12

6
13

b has -CHO

Photosynthesis
• H20 + CO2  (CH2O) + O2
• Chloroplast thylakoid
membranes have chlorophyll
a and b

14

7
 Absorbance of
Chlorophyll a & b
note lack of absorbance 500-600 nm

So photosynthesis
uses mostly blue

!
and red light!

15

Fig. 9.2

Excited
e– state
Energy of electron

Heat

Photon
(fluorescence)
Photon Ground
Chlorophyll state
molecule

(a) Excitation of isolated chlorophyll molecule (b) Fluorescence

16

8
17

Light energy has

now been turned
into chemical
energy!!!!!

Light harvesting systems / antennae

Channel energy through quantum tunnelling
Note: it is not electron tunnelling

18

9
 How a photosystem harvests light

Robert Hill
1899-1991

19

Note that there are two photosystems II and I

20

10
 Photosystem 2

Electron transfer and proton translocation

Pheophytin
chlorophyll minus Mg2+

e e e
Qa Qb
Qb Plastoquinone/ol
Q  QH2
e e

Stroma
PC e
Qb
PC e
Cyt
bf
complex
e e
2 x P680

4H+ in to thylakoid space adds to proton gradient Thylakoid lumen

21

Splitting of water and

O2 release at PSII
H2O Stroma

O There are now

electron “holes”,
Mn Mn Electron “holes”,
i.e. they are
E-
E-
Mn Mn filled
missing

E- E-

2xH Thylakoid lumen

P680

H+ in Thylakoid space - Adds to proton gradient

22

11
 How linear electron flow during the light reactions
generates ATP and NADPH
Please note that the cytochrome complex does not directly
make ATP,
This cartoon is highly misleading!
It does so indirectly, it transfers 4 H+ just like Complex III of
the mitochondria
These then drive ATP synthesis as in the mitochondria

23

Photosystem 1 (P700) Stroma

Acceptor chlorophylls and phyloquinones

e-
Ae- A1Fe-S
-- --0
Fe-S
Fe-S
Ferredoxin
NADP
e- reductase
PC --
NADP+ NADPH
PC e-
--

Cyt
bf
comple e- e-
x -- --
Thylakoid lumen
P700

24

12
 Photosystem 1 (P700) Stroma

e-
Ae- A1Fe-S
-- --0
Fe-S
Fe-S
Ferredoxin
Now there restored
Electrons are NADP
again electron reductase
PC e-
-- holes NADP+ NADPH
PC e-
--

Cyt
bf
complex e- e-
-- --
Thylakoid lumen
P700

25

A tentative model for the organization of the thylakoid membrane

STROMA
(low H+ concentration) Cytochrome
Photosystem II Photosystem I
complex
4 H+ Light NADP+
Light reductase
Fd 3
NADP+ + H+

Pq NADPH

e– Pc
e– 2
H2O
THYLAKOID SPACE 1 1/
2 O2
(high H+ concentration) +2 H+ 4 H+

To
Calvin
Cycle

Thylakoid
membrane ATP
STROMA synthase
ADP
(low H+ concentration)
+ ATP
Pi
H+

26

13
 There is a proton gradient developed by:
1) The splitting of water  H+ released in thylakoid space
2) Cytochrome complex transfers 4 H+ to in thylakoid space
H+
H+
H+ H+ H+ H+
H++ H+ H+
H
2H + O
H+
H2O

NADPH
H+
ATP H+
H+ H
+

Chloroplast

27

Chloroplast ATPase

Is very similar across species, but there are 14 c-ring subunits in plants chloroplasts, only
8 subunits in animal and plant mitochondria
As it turns each subunit binds a proton.
Potential exam question!
What does this mean for ATP synthesis efficiency (ATP/H)?

28

14
 Mitochondrion Chloroplast

MITOCHONDRION CHLOROPLAST
STRUCTURE STRUCTURE
H+ Diffusion
Intermembrane Thylakoid
space space
Electron
Inner Thylakoid
transport
membrane chain membrane

ATP
synthase
Matrix Stroma
Key
ADP + P i
ATP
Higher [H+] H+
Lower [H+]

29

H+
H+
H+ H+
H+ H+
H+ H+
H+ H+
H+
H+

+
-150 m V

CI CIII CIV -
H+
H+

There is another difference in the ATP synthesis capacities

when comparing mitochondria and chloroplasts
There are differences in membrane permeability
Mitochondria  uses the proton gradient and electrical charge

30

15
 In chloroplasts the thylakoid membrane is permeable to Cl-
and Mg2+
H +
H+
H+ H+ H+ H +
H+ H+
H + H+
H + H+ H +
H+
H+
H+ H+ H+
Mg2+

Mg2+ H + Mg 2+

H+ H+
H+

Cl- Cl- H+
Cl- H+
H+ H
+

Chloroplast  proton gradient only, very low pH

31

Differences between mitochondria and

chloroplasts
Chloroplast inner membranes are permeable to Cl- and Mg2+

Chloroplasts are therefore reliant on pH (H+ gradient)

Mitochondria can use net charge potential and H+ gradient

Larger pH difference required in chloroplasts as only H+

gradient is used.

32

16
 Photosystem II and I separated?
Possibly to do with ….
Cyclic Electron Flow
33

Cyclic electron flow

34

17
 Photosystem 1 P700

A0
A1
Fe-S
Ferredoxin

PQ

Cyt
bf
comple
PC
x PC
ee
ee
P700
H+
Cyclic electron flow
35

Why have cyclic electron

flow?
-This produces ATP only (no NADPH)
-Some bacteria only have PSI
36

18
 The plant may simply want more ATP!

ATP is required in C4 plants

What are C4 plants?

….

37

Part 2 Melvin Calvin, a man with mean parents?

Key Concepts (the big picture):

•The light reactions produce O2, H+, ATP and NADPH.

•The Calvin cycle uses NADPH and ATP to fix atmospheric
CO2 to form glyceraldehyde 3-phosphate (G3P).
•Ribulose bisphosphate (RuBP) is used to bind/fix CO2 by the
enzyme RUBISCO (ribulose bisphosphate carboxylase)
•The consumption of ATP and NADPH by the Calvin Cycle is
sustained by the light reactions.

38

19
 WHAT YOU NEED TO KNOW!! You should:

• describe the processes taking place in the Calvin

cycle
• discuss how the absorption of light is coupled to
the synthesis of glucose

39

A simple Calvin Benson cycle

40

20
 The Calvin Cycle is confined to the Stroma

This partitioning stops intermediates and products

(e.g. glyceraldehyde 3 phosphate) being
metabolized, as some of the intermediates feed
directly into glycolysis.

Plants also have mitochondria and glycolytic

enzymes!

41

The Calvin Cycle

This is the level you

need to understand

42

21
 Three Phases of the Calvin
Cycle

1. CO2 fixation
3. Regeneration

2. Reduction

43

Three Phases of Calvin Cycle

1. CO2 fixation

44

22
 Input 3 (Entering one
at a time)
CO2
6C3C
Phase 1: Carbon fixation
5C Rubisco

3 P P
Short-lived
intermediate
3P P 6 P
Ribulose bisphosphate 3-Phosphoglycerate
(RuBP)

3 x 5C  3 x 6C 6 x 3C

45

Ribulose Bisphosphate Carboxylase (RuBisCo)

carboxylates ribulose-1,5-bisphosphate, (5-C
compound) with CO2

x 2!!!

A 2-step reaction gives extremely unstable 6-C

intermediate. This splits in half to form 3-
phosphoglycerate (3PGA), a 3-carbon compound.

46

23
 Fig. 10-18-2
Input 3 (Entering one
at a time)
CO2

Phase 1: Carbon fixation

Rubisco

3 P P
Short-lived
intermediate
3P P 6 P
Ribulose bisphosphate 3-Phosphoglycerate
(RuBP) 6 ATP

6 ADP

Calvin
Cycle
Again note multiple 6 P P
1,3-Bisphosphoglycerate

molecules pass 6 NADPH

through the 6 NADP+

6 Pi

reactions 6 P
Glyceraldehyde-3-phosphate Phase 2:
(G3P) Reduction

1 P Glucose and
Output G3P other organic
(a sugar) compounds

47

Enzymes similar to glycolysis!

Again note multiple molecules pass through the

reactions and therefore 6 x ATP and NADPH
Reduction (gain of electron) and de-
phosphorylation of carbon

48

24
 As a very rough rule NADP/NADPH is involved in reducing
reactions NAD/NADH in oxidation!

NADPH tends to be used for biosynthesis / regeneration

49

Input 3 (Entering one

at a time)
CO2

Phase 1: Carbon fixation

Rubisco

3 P P
Short-lived
intermediate
3P P 6 P
Ribulose bisphosphate 3-Phosphoglycerate
(RuBP) 6 ATP

6 ADP

3 ADP Calvin
Cycle
6 P P
3 ATP
1,3-Bisphosphoglycerate
6 NADPH
Phase 3:
Regeneration of 6 NADP+
the CO2 acceptor 6 Pi
(RuBP)
5 P
G3P
6 P
Glyceraldehyde-3-phosphate Phase 2:
(G3P) Reduction

1 P Glucose and
Output G3P other organic
(a sugar) compounds

50

25
 We will settle with “the regeneration
phase”

After the loss of 1 x C3

5 x C3 are left,
Then these are rearranged to 3 x
C5 (ribulose bisphosphate)
& RuBP is regenerated!

51

The Calvin Cycle takes…

3CO2 6NADPH 9ATP → 6G3P

1G3P is removed/cycle!
So 2 cycles required to make 1 glucose
(The Calvin cycle releases 1 three-carbon sugar phosphate/turn,
A triose phosphate = glyceraldehyde-3-phosphate (G3P)
Hexoses (e.g. glucose) are made using enzymes of gluconeogenesis (coming))
Hexose sugars are not products of Calvin Cycle, C6H12O6, is convenient as it balances
with glycolysis.

52

26
 Thus, only 1 G3P available for subsequent
conversion to ½ a hexose
This required 9 ATPs and 6 NADPH
per 3 CO2 fixed
Energetically $$$$$!
Lucky sunlight is cheap

53

Photorespiration and Issues with Carbon Fixation and Water

The Calvin Cycle is regulated by light in several ways

(The Dark Reactions are a misnomer)

1) pH! The more alkaline the Stroma, the faster the CC

works, the stromal pH rises when the light reactions
are working.
2) Ferrethio reductase (an antioxidant enzyme of sorts)
needs NADPH to reduce/stabilse RuBisCo.
3) Explains why RUBISCO doesn’t work well in isolation
and it needs light

54

27
 RuBisCo has major issues (-or not?)
Rubisco is slow!
It also binds O2 in the process photorespiration
This makes a useless/toxic product (2-phosphoglycolate) and wastes
water!
Also the faster RuBisCo works it makes more errors (high
temperature increases reaction rates & errors!)
When O2 is high (& low CO2) up to 20-30% of photosynthesis
is wasteful! Not so good for farmers!

55

In the Cambrian 33% Oxygen and very

low CO2,
Maybe photorespiration may remove
reactive O2 and this leaves CO2
Still, this must slow growth and requires
more water?

Some plants have evolved a solution

56

28
 C4 plants: Evolution of a solution for photorespiration

Corn is a C4 plant it needs

much less water

oxaloacetate

Rice is a C3 plant
Requires much water

Some plants (hot climates) can concentrate CO2 into

oxaloacetate (4C)

They are called C4 plants (sugarcane and corn).

There is partitioning of specific cycles into different leaf

cells. 4-Carbon oxaloacetate is used as a “CO2” shuttle.

57

C4 plants have different structures in leaves

58

29
 Shuttle enzymes we associate with the
CAC and gluconeogenesis concentrate
CO2 for the CC

Malate and Oxaloacetate have 4 Cs

This acts like a chemical pump, which collects CO2 and concentrates it
This vastly decreases competition with O2 at Rubisco!

59

But why?
If C3 plants photosynthesise at a high rate, O2
accumulates and results in photorespiration.

This is a wasteful use of H2O and the carbon already in the

CC.

The C4 system concentrates CO2 and decreasing for

photorespiration.

But C4 plants uses more ATP!

This extra ATP comes from cyclic electron flow and

sunlight is cheap, especially in the tropics.

Ultimately C4 plants use water more efficiently and

function better in warm climates.

60

30