The light reactions of photosynthesis

Objective of the lecture:

1. To describe the structure of function of chloroplasts.

2. To define the light reactions of photosynthesis.

Text book pages:

198-212.
 Photosynthesis Chapter 10 of text book
Plants use sunlight, carbon dioxide, and water to produce carbohydrate
with oxygen as a byproduct.
The overall chemical reaction summarizes the process as:

CO2 + 2 H2O + light energy  (CH2O)n + H2O + O2

where (CH2O)n stands for carbohydrate.

Usually, glucose (C6H12O6) is considered as the carbohydrate made so:

6 CO2 + 12 H2O + light energy  C6H12O6 + 6 H2O + 6 O2

... this may keep the chemists happy

... but a better summary is of how the process occurs is:

Light-dependent
reactions Calvin cycle
Light Chemical Chemical
energy energy energy

Sunlight H2O O2 ATP, NADPH CO2 (CH2O)n

Thylakoid Reactions Stroma Reactions

Light reactions Dark reactions
 Plant structure, particularly cell structure
(1) makes the reactions possible,
(2) enables integration of light and dark reactions.

Leaves contain millions of chloroplasts.

Cell

Chloroplasts

Fig. 10.2
 Chloroplasts are highly structured, membrane-rich organelles.

Outer membrane
Outer membrane

Inner membrane
Inner membrane

Thylakoids
Thylakoids

Granum
Granum

Stroma
Stroma

Recall that membranes are

composed of a lipid bilayer in
which are embeded proteins
that enable exchange of
materials across the
membrane.

Fig. 6.13
Phospholipids are
in constant lateral Phospholipid
motion, but rarely bilayer
flip to the other
side of the bilayer

Membrane proteins
Figure 6-18b
 There are two processes in photosynthesis that capture light and produce
energy rich compounds that are used in carbon fixation. These are termed
Photosystem I, and
Photosystem II.
These processes are linked in what is termed the Z scheme of photosynthesis.
The Z refers to changes in redox potential of electrons.
Note that PSII comes before PSI in this scheme

Wavelength of maximum
absorption in the far red

Wavelength of maximum
absorption in the red
 Light reactions occur in
the thylakoids (PSII) and
stroma lamella (PSI).

Dark reactions in
occur in the stroma
Thylakoid membranes appear stacked like coins but
in fact are highly folded and have a well defined
interior and exterior with respect to the stroma
Fig. 10.8
Chlorophyll is the most abundant pigment in the chloroplast.
All eukaryotic photosynthetic organisms contain both chlorophyll a
and chlorophyll b
-carotene Carotenoids transfer
energy from photons to
chlorophyll. They also
can quench free radicals
by accepting or stabilizing
unpaired electrons and so
protect chlorophyll
molecules

Chlorophylls a and b
When a photon strikes its energy
can be transferred to an electron
in the “head” region. The
electron is excited, raised to a
higher electron shell, with greater
potential energy

Tail

Ring structure in “head”

(absorbs light)
 Wavelengths (nm)

Gamma Ultra- Micro- Radio

rays X-rays Infrared waves waves
violet

The
electromagnetic
spectrum
Shorter Longer
wavelength wavelength
Visible light
nm

Higher Lower
energy energy
 e– Blue photons excite electrons to
an even higher energy state
Figure 10-9

e– Red photons excite electrons

to a high-energy state

Photons

Energy state of electrons in chlorophyll

Fig. 10.6a Different pigments absorb different wavelengths of light.

Chlorophyll b Chlorophylls absorb blue and red

light and transmit green light
Chlorophyll a
Carotenoids absorb blue
Carotenoids and green light and
transmit yellow, orange,
or red light
Fig. 10.6b Pigments that absorb blue and red photons are the
most effective at triggering photosynthesis.

The oxygen-seeking bacteria

congregate in the wavelengths
of light where the alga is
producing the most oxygen

Oxygen-
seeking
bacteria

O2 O2

Filamentous alga
Basic concept of energy transfer during photosynthesis
 Three Fates for Excited Electrons in Photosynthesis

FLUORESCENCE or RESONANCE or REDUCTION/OXIDATION

Electron drops back down to Energy in electron is transferred to Electron is transferred to

lower energy level; heat and nearby pigment. a new compound.
fluorescence are emitted.

Higher Electron
acceptor
Reaction
center

Photon

Photon Fluorescence
e–
Heat

e– e–

Lower Chlorophyll molecule Chlorophyll molecules in antenna complex Reaction center

Photochemistry
The energy of the excited state causes chemical reactions to
occur. The photochemical reactions of photosynthesis are
among the fastest known chemical reactions. This extreme
speed is necessary for photochemistry to compete with the
other possible reactions of the excited state.
Funneling of excitation from antenna system toward reaction center

The excited-state energy of pigments

increases with distance from the
reaction center. Pigments closer to the
reaction center are lower in energy
than those farther from it. This energy
gradient ensures that excitation
transfer toward the reaction center is
energetically favorable and that
transfer back out to the peripheral
portions of the antenna is energetically
unvavorable.
2-D view of structure of the LHCII antenna complex from higher plants

Stroma

Thylakoid Lumen
 Pheophytin has the structure of
In photosystem II, excited
chlorophyll but without the Mg in
electrons feed an electron
the porphyrin-like ring and acts as
transport chain.
an electron acceptor.

Higher
Pheophytin

e–
PQ
Cytochrome
complex

Photon
2. Electrons that reach pheophytin are transferred to
plastoquinone (PQ), which is lipid soluble, passed to
an electron transport chain (quinones and
cytochromes)
Chlorophyll

Lower
1. When an electron in the reaction center chlorophyll
2H2O O2+ 4H+ + 4e- is excited energetically the electron binds to pheophytin
and the reaction center chlorophyll is oxidized
Photosystem II Feeds an ETC that Pumps Protons
Plastoquinone carries protons to
the inside of thylakoids, creating
a proton-motive force.
3. Passage of electrons along the chain
involves a series of reduction-oxidation
reactions that results in protons being pumped
from stroma to thylakoid lumen

Stroma
Stroma Photon The ph of the lumen reaches 5
Photosystem II H+ Cytochrome
Antenna complex while that of the stroma is
complex
around 8 - the concentration of
e– H+ is 1000 times higher in the
PQ
Pheophytin lumen than the stroma.
e–
e– An essential component of the
Reaction PQ
reaction is the physical transfer
center
of the electron from the excited
H2O
chlorophyll. The transfer takes
O2+ H+ H+ H+
Thylakoid Lumen H + H+ ~200 picoseconds (1 picosecond
H+
H+
(low pH) H+ + +
H+
H+ = 10-12 s).
H H

The oxidized reaction center of the chlorophyll that donated an electron is re-reduced by a
secondary donor and the ultimate donor is water and oxygen is produced.
Figure 10-14
Higher Photosystem I

2e–
NADP+ + H+

Iron and sulphur Ferredoxin

compounds

NADP reductase NADPH

2 Photons NADPH is an electron carrier that
can donate electrons to other
compounds and so reduce them.

Chlorophyll

Lower
 The Z scheme linking Photosystem II and Photosystem I

Fig. 10.15
4e–

Higher 2 NADP+ + 2 H+

Pheophytin
Ferredoxin
4e–
PQ
Cytochrome 4 Photons 2 NADPH
complex

4 Photons PC
ATP
produced via P700
proton-motive force Photosystem I
P680
Photosystem II

Lower 4e–
2 H2O 4 H+ + O 2

When electrons reach the end of the Photosystem II electron

chain they are passed to a protein plastocyanin that can diffuse
through the lumen of the thylakoid and donate electrons to
Photosystem I. Shuttle rate of 1000 electrons per second
between photosystems.
ATP synthase – only in the stroma lamella and edge of grana stacks
Stroma Chemiosmosis
Ion concentration differences
and electric potential
differences across
membranes are a source of
energy that can be utilized
Hydrophilic
As a result of the light
reactions the stroma has
become more alkaline (fewer
H+ ions) and the lumen more
acid (more H+ ions)

The internal stalk and much

Hydrophobic of the enzyme complex
located in the membrane
rotates during catalysis.

Thylakoid Lumen The enzyme is actually a

tiny molecular motor
T
Transfer of electrons and protons in the thylakoid membrane is carried out vectorially

Stroma

Thylakoid Lumen
Protons diffuse to the site of ATP synthase

Dashed lines represent electron transfer

Solid lines represent proton movement

Organization and structure of the four major protein complexes

Stroma

LHCI, PSI, and ATP

synthase are all in the
stroma lamella or on the
edge of a grana

LHC light harvesting complex

 Organization and structure of the four major protein complexes

Stroma

Thylakoid Lumen
 Things you need to know ...

1. The structure of chloroplasts and how the light reactions are

distributed and supply ATP and NADPH to the dark reactions

2. The Z scheme of photosynthesis, its photochemical and electro-

potential characteristics and its spatial arrangement
through the chloroplast membrane system, acidification of
the thylakoid lumen and formation of ATP.

3. The energy transfer system during photosynthesis including the

role of different pigments, the antenna and reaction center