Photosynthesis – the electron transport chain II

HBTBC/HBC 206 Plant Biochemistry

L4
 Outline
1. The electron transport chain in plants
2. ATP is generated by chemiosmosis
3. The electron transport chain in purple bacteria
4. The noncyclic electron transport chain in
green sulphur bacteria
5. Two photosystems can create more energy
than the sum of its single photosystems
6. Cyclic vs non-cyclic electron transport
7. Evolutionary origin of the reaction centre’s
of photosystems I and II
The electron transport chain in
purple bacteria
 The electron transport chain in
purple bacteria
• Bacteria are useful for studying the principles of
photosynthesis because their reaction centers
are simpler than those of plants

• They are also more easily isolated

 The electron transport chain in
purple bacteria
Purple bacteria have only one reaction centre
The reaction centre of purple bacteria
The reaction center is a transmembrane protein–
pigment complex P870
The reaction centre of purple bacteria
There are 3 hydrophobic protein subunits
Blue P870

Purple
Orange
The reaction centre of purple bacteria
There are
4 Bacterial Chlorophyll b
2 Bacterial Pheophytin b
FE II ion P870

Ubiquinone
Menaquinone
Converting light to
chemical energy
• A photon is absorbed by the
two chlorophyll molecules at
the top
• An electron is then passed
through the reaction centre
• Ubiquinone is the final
electron acceptor within the
reaction centre
 Reaction Centre excitation sequence
A photon is absorbed by the “special pair” of BChl
b molecules, raising them to an excited state
 Reaction Centre excitation sequence
An excited electron is passed to the BPheo b of
the L subunit
The special pair is left with a positive charge
 Reaction Centre excitation sequence
The excited electron is transferred to the
menaquinone
 Reaction Centre excitation sequence
• The special pair is reduced by a complex electron
transport chain eliminating its positive charge
• The excited electron migrates to the ubiquinone
 Reaction Centre excitation sequence
After a second electron is transferred to
ubiquinone, it picks up two protons from solution
and exchanges with the membrane-bound
ubiquinone pool
 The electron transport chain in
purple bacteria
• The ubiquinone carries the electrons to a
cytochrome bc1 complex embedded in the
membrane
• The electrons are then transferred to a smaller
mobile cytochrome complex that transfers the
electrons back to the reaction centre
• Becaue the excited electrons are returned to the
ground state of the reaction center, this electron
transport is called cyclic electron transport
The cyclic electron transport chain in
purple bacteria
The cyclic electron transport chain in
purple bacteria
 Reduction potential
• The vertical axis in the figure represents the
reduction potential of a particular species
• The higher the position of a molecular species,
the more negative its reduction potential, and
the more easily it donates electrons
 The cyclic electron transport chain in
purple bacteria
***NADH is produced using the H+ membrane potential
The noncyclic electron transport
chain in green sulphur bacteria
 The noncyclic electron transport
chain in green sulphur bacteria
• Green sulphur bacteria are able to form NADH
by a noncyclic electron transport process
• The excited electrons are transferred to the
ferredoxin-NAD-reductase complex
• This complex reduces NAD to NADH
The noncyclic electron transport chain
in green sulphur bacteria
 The noncyclic electron transport
chain in green sulphur bacteria
• The excited electrons in this noncyclic
pathway do not return to the ground state
• Therefore the reaction center remains
oxidised and must be reduced by another
electron donor
• H2S can be oxidized to sulphate in green
sulphur bacteria
The noncyclic electron transport chain
in green sulphur bacteria
 Cyclic electron transport in green
sulphur bacteria
• Green sulphur bacteria can also carry out cyclic
electron transport but this will not produce NADH
• Cyclic electron transport does contribute to the
H+ potential
Cyclic and noncyclic electron
transport chains
 Photosynthesis in plants and
cyanobacteria generates O2
• Neither of these bacterial systems generates
sufficient oxidizing power to oxidize H2O
• They are both anoxygenic and anaerobic
• Compare the oxidising power of the
photosystems of bacteria with those of plants -
Electron transport in different
photosystems
 Two photosystems working together in
noncyclic photophosphorylation enhance the
rate of photosynthesis

Experiments measured
the rate of
photosynthesis using two
light beams of different
wavelengths: one red
and the other far-red
 The two photosystems work together
in noncyclic photophosphorylation
Using both beams produced a rate greater than
the sum of the rates using individual beams of
these wavelengths
 The two photosystems work together
in noncyclic photophosphorylation
• This is called the enhancement effect
• It can be explained by a mechanism involving
two photosystems acting in series (that is, one
after the other)
– one photosystem absorbs preferentially in the red
– the other in the far-red
 Noncyclic photophosphorylation
• The passage of an electron pair from water to
NADPH in noncyclic photophosphorylation
generates one molecule of NADPH and slightly
more than one molecule of ATP.
• Building organic molecules takes more energy
than that—it takes 1.5 ATP molecules per
NADPH molecule to fix carbon.
 Cyclic Electron Flow
• To produce the extra ATP, many plant species
are capable of short-circuiting photosystem I

• This switches photosynthesis into a

cyclic photophosphorylation mode

• The light-excited electron leaving photosystem I

is used to make ATP instead of NADPH.

• The relative proportions of cyclic and noncyclic

photophosphorylation in these plants
determine the relative amounts of ATP and
NADPH available for building organic molecules
 Cyclic Electron Flow
• Cyclic electron flow uses only photosystem I and
produces ATP, but not NADPH
• No oxygen is released
• Cyclic electron flow generates surplus ATP, satisfying
the higher demand in the Calvin cycle
Primary
Primary acceptor
Fd
acceptor Fd
Pq NADP
NADP + H
Cytochrome reductase
NADPH
complex
Notice that this is very
Pc similar to the electron
transport chain in
green sulphur bacteria
Photosystem I
that can carry out both
Photosystem II ATP cyclic and non-cyclic
electron transport.
 Cyclic Electron Flow in PSI
• Notice that this is very similar to the electron transport
chain in green sulphur bacteria
• Green suphur bacteria can also carry out both cyclic
(produces ATP) and non-cyclic electron transport
(produces NADH)
Primary NADP
Fd acceptor NADP + H
Fd ductase
NADPH
q

Cytochrome re
complex
Pc

Green sulphur bacteria

Photosystem I
ATP
Electron transport in different
photosystems
 Evolutionary origin of the
photosystems
• The reaction centre of PS II in plants and
cyanobacteria is a Q (quinone) type RC similar to
that in purple bacteria
• The evolutionary origin of PS II is likely to be
purple bacteria
• The reaction centre of PS I is an FeS type RC
similar to that in green sulphur bacteria.
• PS I can carry out cyclic and non-cyclic electron
transport similar to that in green sulphur bacteria
• The evolutionary origin of PS I is likely to be green
sulphur bacteria
Electron transport in different
photosystems
Summary
1. The electron transport chain in purple
bacteria is cyclic and has a quinone (Q)
type reaction centre
2. Green sulphur bacteria can carry out non-
cyclic (produces NADH) and cyclic
(produces ATP) electron transport
3. Green suphur bacteria have an iron-sulphur
(FeS) type reaction centre
Summary
4. Two photosystems working together in
plants and cyanobacteria can create more
energy than the sum of its single
photosystems
5. This enables oxygenic photosynthesis
Summary
6. PS I can carry out non cyclic (produces
NADPH) and cyclic (produces ATP) electron
transport
7. The evolutionary origin of the reaction
centre’s of photosystems I and II are likely
green sulphur bacteria and purple bacteria
respectively