Module 3: Photosynthesis

Introduction
Photosynthesis literally means “synthesis using light." Life on earth ultimately depends on energy derived
from the sun. Photosynthesis is the only process of biological importance that can harvest this energy.
▪ The process of synthesis of organic food (carbohydrates) from CO2 and water in the presence of light
by plants - photosynthesis
▪ Conversion of light energy into
chemical energy that is stored in
organic compounds. Light energy
drives the synthesis of
carbohydrates from CO2 (C fixation)
and water with the generation of
oxygen.
▪ Plants store energy in the chemical
bonds of sugars. Energy stored in
these molecules can be used later to
power cellular processes in the plant
and can serve as the energy source for all forms of life. Chemical energy is released as ATP during
cellular respiration. Plants use glucose as food for energy and as a building block for growth.
▪ Autotrophs make glucose and heterotrophs are consumers of it. Almost all plants are photosynthetic
autotrophs, as are some bacteria and protists.

An overview of photosynthesis
• Photosynthesis is a chemical, oxidation-reduction process
by which autotrophic organisms use light energy to make
sugar and oxygen gas from carbon dioxide and water.
Plants use solar energy to oxidize water, thereby releasing
the oxygen and to reduce CO2, thereby forming large
carbon compounds (sugars).
• The mechanism of photosynthesis can be represented by the following equation.

• The reduction of CO2 into carbohydrate requires assimilatory powers like ATP and NADPH
• The reduction of CO2 takes place in light independent conditions while the production of assimilatory
powers takes place in presence of light.

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2 phases of photosynthesis
1) Light dependent phase – it requires light and is also called photochemical reaction or light
reaction or Hill –reaction
2) Light independent phase – it does not require light and is also called bio-chemical dark reaction or
dark reaction or Blackman’s reaction
The electromagnetic spectrum

Photosynthetic pigments absorb the light rays with

Wavelength between 400 and 700 nm.

Hence light between 400nm and 700nm is called

Photosynthetically Active Radiation (PAR)

Above 680 nm - far red

Below 680 nm - short red

Dual nature of light

Particle and wave nature of light- light behaves as if it were composed
of "units" or "packets" of energy (quantum) that travel in waves. These
packets of light energy are called photons. The wavelength of light
determines its color.

Light reaction and Dark reaction

In the light reactions, excited electrons (from the photosynthetic pigment - chlorophyll) are moved among a
series of electron carriers that are embedded in the thylakoid membrane of the chloroplasts. The energy
obtained by this electron transport drives the synthesis of ATP in the stroma, and the final step in the light
reactions is formation of NADPH. In the dark reactions, the ATP and NADPH are used to convert CO 2 to
carbohydrate. The dark reactions begin in stroma.

The first stage of photosynthesis, the light reactions, uses the energy of light to reduce NADP (an electron
carrier molecule) to NADPH and to manufacture ATP. The NADPH and ATP from the first stage of
photosynthesis are then used in the second stage (dark reaction), the Calvin cycle, to reduce the carbon in
carbon dioxide and form a simple sugar.

Chloroplast – The site for Photosynthesis

In most plants, photosynthesis occurs primarily in the leaves. The most active photosynthetic tissue in
higher plants is mesophyll of leaves. Mesophyll cells have many chloroplasts (green plastids seen in
cytoplasm- membrane bound organelle), which contain the specialized light absorbing green pigments-
chlorophylls.

Chloroplasts are homogenously distributed in the cytoplasm. Their diameter is about 5 to 8 microns. Each
mesophyll cell may contain as many as 300 chloroplasts.

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They have various shapes in green algae - cup
shaped in Chlamydomonas, Spiral in
Spirogyra, star shaped in Zygnema etc. The
chloroplast is surrounded by 2 unit
membranes. Each membrane is lipo-
proteinaceous in nature. The space between 2
membranes is called as periplastidal space.

A chloroplast contains:
1. An outer membrane - highly permeable
2. An inner membrane - nearly impermeable
3. Stroma - proteinaceous fluid matrix filling the space inside the chloroplast. It is aqueous and contains
enzymes for carbon assimilation, DNA, RNA, ribosomes. The enzymes found in stroma are capable of
utilizing ATP and NADPH to produce carbohydrates.
4. Grana - most of the thylakoids appear to be very closely associated with each other. Such stacks of
thylakoids are called grana/ grana lamellae. Exposed membrane in which stacking is absent is called as
stroma lamellae. Grana lamellae are connected by “stromal lamellae”

Thylakoids/lamellae - membrane bound sac like structures (a membranous compartment) embedded in the
stroma. It is derived from invaginations of the inner membrane. The thylakoids contain chlorophyll.

Structure of chloroplast is very similar to mitochondrion and probably evolved from a cyanobacterium
incorporated into a non-photosynthetic eukaryote (endosymbiosis). In eukaryotes, the light reaction occurs
in thylakoid membrane and the dark reaction occurs in the stroma. While in prokaryotes, the light reaction
occurs in the invaginations of inner (plasma) membrane (by embedded chromatophores)

Photosynthetic pigments
During photosynthesis light energy must be absorbed by some pigments. A pigment is a molecule that
absorbs light of a specific wavelength in the visible spectrum. Photosynthetic pigments seen as groups in
the thylakoids are called- photosynthetic units.

Chloroplast contains several major pigments

1. Chlorophyll a
2. Chlorophyll b
3. Carotene
4. Xanthophyll
5. Phycobilins

2 major types of chlorophyll

Chlorophyll is the green photosynthetic pigment. There are different types of chlorophyll - Chl a, Chl b,
Chl c, Chl d, Chl e, bacteriochlorophyll etc. Of all Chl a & b are widely distributed in green algae and
higher plants.

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The basic structure
▪ In porphyrin head, 4 pyrrole (tetrapyrrole) rings are linked together by methane groups to form a
ring system. The center of tetrapyrrole is occupied by a central core of magnesium (Mg). 2
covalent and 2 co-ordinate bonds bind the Mg atom with the pyrrole ring.

▪ A lengthy hydrocarbon tail or side chain is attached to the porphyrin ring- phytol.
▪ Chlorophyll has a tadpole like structure. Porphyrin head is hydrophilic and Phytol tail is
hydrophobic
▪ Chlorophyll a – involved in light reactions
▪ Chlorophyll b – assists in capturing light energy – accessory pigment
▪ Chl a and Chl b structurally differs in the nature of groups attached. In Chl a, a methyl group (-CH3)
and in Chl b, an aldehyde group (-CHO) are attached to the 3rd carbon of the 2nd pyrrole ring in the
porphyrin head. CHO group in Chl b increases the blue light absorption and decreases the red light
absorption peak. Both are soluble in organic solvents like alcohol, acetone etc.

▪ Chlorophyll molecules absorb only red, blue and violet lights in the visible spectrum. They reflect
green light. The maximum rate of photosynthesis has been observed in red light.

▪ Chl a is known as universal photosynthetic pigment. The photochemical reaction (light reaction)
takes place only in Chl a.

▪ Chla molecules are of different types. Chl a 673, Chl a 683, Chl a 680, Chl a 700.

Difference between Chl a and Chl b

Chl a Chl b
It is a primary photosynthetic pigment. It is an accessory pigment
Molecular weight - 873 g/mol Molecular weight - 907 g/mol
Blue green in pure state Olive green in pure state
Empirical formula is C55H72O5N4Mg Empirical formula is C55H70O6N4Mg
rd rd
In Chl a, a methyl group (-CH3) at 3 carbon of In Chl b, an aldehyde group (-CHO) at 3
pyrrole ring. carbon of pyrrole ring.
More soluble in petroleum ether More soluble in 90% methyl alcohol.
It occurs in all photosynthetic organisms except It occurs in all green plants except diatoms,
photosynthetic bacteria. cyanobacteria, brown algae and red algae.
Carotenoids
They are yellow, brown, orange or red accessory pigments found in close association with chlorophyll in
chloroplast. They capture more light energy and absorb blue and green lights in the visible spectrum.

Functions:-
1. Absorb light energy and transmit it to the neighbouring pigment molecules in the photosynthetic unit.
2. Protect the chlorophyll molecules from photo-oxidation by picking up nascent O2 and converting it
into harmless molecular stage.

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In general, there are 2 groups of carotenoids.

1. Carotenes (orange pigment) - general formula is C40H56

They are abundant in carrot, tomato etc. Important carotenes are Beta carotenes (β carotenes – mole.
weight -536.873 g/mol).

2. Xanthophyll (yellow pigment) - general formula is C40H56O2.

They are more abundant than carotenes. Found in autumn leaves. Mole. weight - 568.9 g/mol.

Phycobilin
It comprises a major group of photosynthetic accessory pigments in algae
▪ Phycocyanin (blue colour) – in blue green algae
▪ Phycoerythrin (red colour) – in Rhodophycean members.

Mechanism of light absorption and emission

During absorption of light, chlorophyll in its lowest energy or ground state absorbs a photon and makes a
transition to a higher energy or excited state.
Excited chlorophyll has 4 alternative pathways for disposing of its available energy.
1. Fluorescence (a slower process)
Excited Chl can re-emit a photon and thereby return to its ground state – fluorescence. Wavelength of
fluorescence is slightly longer than the wavelength of absorption.

2. Internal conversion
Excited Chl can return to its ground state by directly converting its excitation energy into heat, with no
emission of a photon.

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3. Exciton transfer (“resonance” transfer)
Chl may participate in energy transfer, during which an excited chlorophyll transfers its energy to another
unexcited molecule.

4. Photochemistry (photo-oxidation)
In which the energy of the excited state causes chemical reactions to occur. The excited molecule transfers
its electron to an acceptor molecule. The photochemical reactions of photosynthesis are among the fastest
known chemical reactions. The extreme speed is necessary for photochemistry to compete with the 3 other
possible reactions.

Photoluminescence: fluorescence and phosphorescence

Fluorescence and phosphorescence are two forms of photoluminescence. Photoluminescence is the
emission of light arising from excited electronic states following absorption of light.

Phosphorescence is a specific type of photoluminescence related to fluorescence. In simple terms

phosphorescence is a process in which energy absorbed by a substance is released relatively slowly in the
form of light. Phosphorescent materials “store/trap” electrons in a higher energy state for a long time
(minutes or even hours).

Photosynthetic unit/ Photosystem/ Pigment system

The smallest group of pigment molecules participating in a photochemical reaction is called Photosynthetic
unit/Photosystem. Each Photosynthetic unit has

1. Reaction center complex

2. Light harvesting antennas/ Light Harvesting Complex (LHC)

1. Reaction center complex

It consists of a dimer of special Chl a molecule which absorbs long wave light energy (low energy). This
reaction center chlorophyll plus associated proteins and redox carriers (primary electron acceptor) are
directly involved in light-driven redox reactions.

It is also called Photocentre or Trap centre. Only Chl a gives excited electron to an acceptor.
3 different types of reaction centres
• P700 (in PS I)- absorption peak around 700nm.
• P680 (in PS II)- absorption peak around 680 nm.
• P870 – bacteriochlorophyll with an absorption peak around 870nm.

2. Light Harvesting Complex (LHC)

The reaction center is surrounded by a number of light harvesting pigment molecules. The photocenter
requires the help of light harvesting pigment molecules in the absorption of light energy. Light harvesting
pigment molecules absorb light energy of different wavelengths and transfer it to the neighbouring pigment
molecules (resonance transfer). Finally, the light energy is focussed on to the reaction center. They absorb
those wavelengths of light which is not absorbed by Chlorophyll a. It is important because they absorb
excessive light and protects Chl a.
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LHC - composition
The core antenna chlorophyll a
• These are antenna pigment molecules lie around the reaction centre.
• Their number vary from 20 – 30. They are usually Chl a molecules.
• The core antenna chlorophyll a absorb light but do not participate directly in photochemical
reactions.
Antenna pigment molecules
• Antenna pigment molecules lie around these core antenna Chl a molecules.
• They are usually 200-300 in number. They absorb light energy of different wavelengths but shorter
than that of photocenter.

• E. g. various chlorophyll molecules, carotenoid molecules.

• The antenna system of different classes of photosynthetic organisms are remarkably varied, in
contrast to the reaction centers, which appear to be similar in even distantly related organisms.

• The sequence of pigments within the antenna that funnel absorbed energy towards the reaction
center has absorption maxima that are progressively shifted toward longer red wavelengths.

Absorption spectrum
An absorption spectrum is a graphic representation which displays the amount of light energy take up or
absorbed by a pigment molecule or substance as a function of the wavelength of the light.
E. g. Chl a absorbs blue and red lights in the visible spectrum. The amount of light absorbed by a pigment
can be plotted here.
Action spectrum
Photosynthesis takes place in visible light ranging from 400 nm to 700 nm wavelength. It does not takes
place at the same rate with all wavelengths.

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 • An action spectrum depicts the magnitude of a response of a biological system to light, as a
function of wavelength.

• It is the graphic representation of rate of photosynthesis against the wavelength (1st action spectrum
was constructed for an alga - Spirogyra)

• An action spectrum for photosynthesis can be constructed from measurements of oxygen evolution
at different wavelengths.

• Action spectra were very important for the discovery of 2 distinct photosystems operating in
oxygen evolving photosynthetic organisms.

Red drop and Emerson’s enhancement effect

Quantum yield of photosynthesis –
Number of photochemical products (O2 molecules) released
Q = -----------------------------------------------------------------------------------
Light quanta absorbed
• Robert Emerson and Lewis using monochromatic light (light with specific wavelength/colour)
performed a number of experiments in Chlorella culture to measure the quantum yield of
photosynthesis.
• Quantum requirement is defined as the no. of light quanta required to release one molecule of O2 in
photosynthesis.
• Emerson showed that reduction of one molecule of CO2 to C6H12O6 and liberation of one molecule O2
requires a minimum of 8 quanta of light. Thus, the quantum requirement of photosynthesis is 8 quanta
and quantum yield is 1/8 or 12%.
• When quantum yield was determined under different wavelengths of light, the yield was almost
constant in all the wavelengths but dropped suddenly in the region above 680nm. Although such light
was still absorbed by Chl a. This decrease was noticed in the red zone- red drop phenomenon.
• This pronounced decrease/drop in the quantum yield at wavelengths greater than 680nm (in far
red region) is called red drop phenomenon.
or
• The deficiency in the ability of far-red light to assist photosynthesis – red drop phenomenon.

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 • When repeated the 1st experiment by
supplying additional shorter
wavelength of light along with far red
light it was observed that quantum yield
was enhanced by combined effect of
short and long wavelength of light
beyond 680 nm.

• So, Emerson and co-workers found that

the decrease in quantum yield could be
overcome if shorter wavelength was
given to culture simultaneously.

• Thus, photosynthesis obtained from the

2 combined beams of light was found to
be much higher than the average of the
sum of the production under the
separate beams of light.

• This increase in in quantum yield by

simultaneous application of shorter and
longer wavelengths of light is called
Emerson enhancement.

• These experiments gave conclusive proof that photosynthesis involves 2 photochemical reactions.

• One carried by shorter wavelength and the other by long wavelength of light and driven by 2 groups
of photosystems. Photosystems I and II (PS I & PS II)

Photosystem/ Pigment system (PS)

Photosystem is a complex of pigments and proteins arranged on the thylakoid membrane as functional unit.
The combination of light harvesting complex (LHC) (accessory pigments) and reaction center is called
photosystem.

• Two photosystems
– Photosystem I (P700)
– Photosystem II (P680)

Long red wavelengths are absorbed by PS I. Short red wavelengths are absorbed by PS II. For maximum
photosynthesis, both systems must function together. About 250 to 450 pigment molecules constitute a
single photosystem. Photosystem I evolved very early, and it is found in non-oxygenic phototrophs;
photosystem II evolved later.

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Photosystem I

• Here the reaction centre is P700.

• Here P stands for ‘pigments’.

• The P700 is a type of Chl a molecule with an absorption peak around 700 nm.

• Photochemical reaction takes place only in P700.

• P700 is surrounded by other Chl a molecules, followed by Chl b and carotenoids.

• PS I is located in stroma thylakoids.

• PS I participate in both cyclic and non-cyclic photophosphorylation independently.

• The purple bacteria utilize only one photosystem (PSI).

Photosystem II

• Here the reaction centre is P680.

• The P680 is a type of Chl a molecule with an absorption peak around 680 nm. PSII photosystem is
most sensitive to shorter wavelength 680 nm light, it absorbs slightly more energy than the P700-
PSI system.

• Photochemical reaction takes place only in P680.P680 is surrounded by other Chl a molecules,
followed by Chl b and carotenoids.

• The carotenoid content is higher as compared to PS I.PS II is located in grana thylakoids.

• PS II participates in only non-cyclic photophosphorylation. It can operate only with the help of PS
I.
• Oxygenic phototrophs utilize two photosystems (PSI and PSII).
• Primary electron acceptor in PS II is a colourless Chl a, that lacks Mg2+ - phaeophytin.
Functions of photosystems
• To trap light energy.

• To convert absorbed light energy into chemical energy. This chemical energy is stored in the form
of ATP and NADPH.

2 phases of photosynthesis
STAGE 1 - LIGHT REACTION - energy from sun is captured used to split water into H+ an O2.

▪ Water is Split into Hydrogen Ions, Electrons, and Oxygen (O2).The O2 Diffuses out of the
Chloroplasts (Byproduct).

▪ The Light Energy is Converted to Chemical Energy, which is Temporarily Stored in ATP and
NADPH in stroma.

STAGE 2 – DARK REACTION - where carbon is fixed into glucose.

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 ▪ The Chemical Energy Stored in ATP and NADPH powers the formation of organic compounds
(Sugars), using Carbon Dioxide, CO2.

Steps in light reaction in brief

• Light hits reaction centers of chlorophyll, found in chloroplasts
• Chlorophyll get excited and subsequently causes water to break apart.
• Oxygen is released into air
• Hydrogen remains in chloroplast gets attached to NADP+ to form NADPH
• Formation of ATP
• “THE LIGHT REACTION” (Hill reaction)

Steps in Dark reaction in brief

• The DARK Reactions = Calvin Cycle.

• CO2 from atmosphere is joined to H from water molecules (NADPH) to form glucose.

Light reaction- mechanism

• Photo-excitation of Chlorophyll
• Photolysis or photo-oxidation of water (in non-cyclic photophosphorylation only)
• Photoreduction
• Photophosphorylation
– Cyclic
– Noncyclic
Photo excitation of Chlorophyll

• The first step in photosynthesis is the absorption of light by pigment molecules.

• The P700 and P680 are the reaction centres of photosystem I and photosystem II respectively.

• The photochemical reaction takes place only in P700 and P680.

• When light energy absorbed, they get excited and high energy electrons are soon emitted from it.

Photo-oxidation

• Excited chlorophyll is the donor of electron in photosynthesis.

• After the transfer, chlorophyll is oxidized to a cationic free radical.
• It returns to its ground state by oxidizing another molecule in the case of PS II (P 680).
• P 680 becomes a strong oxidising agent and splits a molecule of water to release oxygen.
• This light dependent splitting of water molecule into hydrogen ion (H+), electrons (e-) and oxygen
(O2) is called photolysis of water.
• Manganese, calcium and chloride ions play important roles in the photolysis of water. It was first
demonstrated by Robert Hill in 1937.

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 • The energy in the electron of the excited chlorophyll of PSII (P680) will be received by its primary
electron acceptor called pheophytin.
• Loss of electron causes P680 to become positively charged (P680+) and it is then attracts an
electron from an adjacent Manganese protein (Mn2+ protein). It acts as water splitting protein
system.
• Mn2+ in turn strongly attracts an electron from H2O.
• Each water can give up only two electrons (Which are covalently held with H2).
• Hence water photolysis yields,

H2O 2 H++ 1/2 O2 + 2e-

or

2 H2O 4 H++ O2 + 4e-

• The consecutive excitation of P680 makes ultimately lose both electrons from water, one at time.

Photo-reduction

• Electron derived from pheophytin (Primary electron acceptor) will move down hill through many
electron acceptors.
• First it will enter into plasto-quinone pool (PQ).
• The PQ thus get reduced. A complete reduction of PQ to PQH2 take place as it receives 2 H+from
stroma side.
• From PQH2 electron move one at a time to cytochrome b6 and then to the iron sulphur proteins (Fe-
S) and later to cytochrome f in the complex between PSII and PSI.
• The 2 H+however will be released to proton reservoir.
• From here e- go to Plastocyanin (PC), which is thought to move along with the edge of thylakoid
membrane to PSI.
• P700 accept this e-.

Light Reaction - Electron Flow

▪ Occurs in the Thylakoid membranes

▪ 2 possible routes for electron flow:

• Use Photosystem I and Electron Transport Chain (ETC) and generate ATP only- CYCLIC

OR

• Use Photosystem II and Photosystem I with ETC and generate O2, ATP and NADPH - NON-
CYCLIC

In the light reactions, electron transport chains generate ATP, NADPH, & O2

• The excited electrons are passed from the primary electron acceptor to electron transport chains.

– Their energy ends up in ATP and NADPH.

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Phosphorylation
• The FREE PHOSPHATE can be re-attached to ADP reforming ATP. Process called
Phosphorylation.
• The addition of an inorganic phosphate to ADP, under the influence of light during the light
reaction of photosynthesis is called photophosphorylation.

• Photophosphorylation takes place by CHEMIOSMOSIS.

• The production of ATP using the energy of hydrogen ion gradient across the membrane to
photophoshorylate ADP is called chemiosmosis.

Cyclic Electron Transport Chain

• Electron transport does not only occur from water to NADP (non-cyclic), but there is also a second
mode of electron transport which involves neither oxidation of water nor reduction of NADP and
only requires excitation of PS I.
• This pathway is called cyclic electron transport because electron cycles from P700 through several
carriers back to P 700.

• Here, absorption of 2 photons activates P 700 and cause 2 electrons to cycle and deposit 2 H+ in the
thylakoid channel when PQH2 is oxidized (translocation of protons across the thylakoid to the inner
side).

• No water is split here and no NADPH is formed.

• But ATP is produced by the coupling factor (CF) in response to the decreased pH in the channel
(more H+ inside thylakoid; less H+ in stroma) causing proton motive force necessary to supply
energy for the phosphorylation of ADP with Pi, the formation of ATP by this cyclic electron
transport pathway, is therefore called cyclic photophosphorylation.

• Non – cyclic electron flow is the normal process occurring in photosynthesis. While cyclic electron
flow involves only the synthesis of ATP.

• cyclic electron flow:

• Independent of PSII
• No o2 evolved, no NADPH formed.

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Non-cyclic Electron Transport Chain
• Non-cyclic electron transport (Known as Z scheme as it resembles English letter ‘Z’) involves the
flow of electrons from water, through PS II and PS I in series to the terminal electron acceptor
NADP. This pathway is called non-cyclic because electrons driven from H2O to NADP never cycle
back.
• When a photon is absorbed by a pigment in the LHC associated with PS II, its energy is transferred
to P680.
• This excites an electron in P680 so much that it can be removed by Pheo.
• Loss of the electron causes P 680 to become positively charged. Which then attracts an electron
from an adjacent Mn-protein. As the Mn-protein loses an electron and becomes oxidized, it then
attracts an electron from H2O.
• Each H2O molecule can give up 2 electrons.
• Thus, as H2O is oxidized by losing 2 electrons, it also forms ½ O2 and 2 H+.
• This oxidation requires the absorption of 2 photons by PS II and 2 consecutive excitations of P680
with loss of 2 electrons, one at a time.
• On the reducing side of PS II, the first electron acceptor is plastoquinone (PQ) and its reduction to
PQH2 requires 2 electrons and 2 H+.
• From there, electrons move to the cytochrome b6 f complex and from which electrons go to
plastocyanin (PC), a copper- containing protein which is the electron donor to P 700(PS I).
• However, P 700 cannot accept an electron unless it has previously loss one and such loss can occur
only by light excitation.
• The reducing side of PS I is rich in Fe-S proteins.
• When electrons pass through a soluble Fe-S protein ferredoxin (Fd), its Fe3+ get reduced to Fe 2+.
• Terminal electron acceptor is NADP and Fd reduces NADP by giving 2 electrons, one at a time to
NADP. This reaction takes place in stroma catalysed by Fd-NADP reductase.
• Water is the ultimate source of electrons delivered to NADP in plant oxygenic photosynthesis.
• Being excited by photon absorption, P680 of PS II loses an electron to phaeophytin and must get an
electron to return to the ground state for the capture of another photon.
• Oxidation of water to form molecular oxygen by chemical means is a difficult task and is achieved
in nature only by the photosynthetic apparatus.
• The reaction centre must be excited 4 times in order to produce a single O2 molecule by the
oxidation of 2 molecules of water.
• Formation of ATP by this electron transport is called non-cyclic photophosphorylation.
• In the ETC, reduction of PQ to PQH2 by Q requires 2 e- and 2 H+.
• PS I and PS II cooperatively function to transfer electrons from H2O to NADP.
• 2 mobile electron carriers carry electrons from PS II to PS I, thus providing the necessary
connection.
• One carrier is a small copper containing protein- plastocyanin.
• When its copper becomes reduced from Cu 2+ (cupric form) to Cu 1+ (cuprous form) by PS II.
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 • It can move along the membrane, carrying an electron.
• Photophosphorylation depends on absorption of these ions (H+) from the stroma side of membrane
as PQ reduction occurs.
• It also depends on the transport of these same H+ ions into the thylakoid channel when PQH2 is
reoxidizedbyCyt f which is the next member in the ETC.
• Here, these H+ combine with those released from H2O oxidation and jointly they cause the pH to
decrease in the channel leading to the build-up of a trans membrane proton concentration
gradient.
• The other carrier system is nothing but a group of quinones, - plastoquinones (PQ) which carry 2
electrons and 2 H+ from PS II to PS I.
• Other electron transport components are 2 cytochromes- cytochrome b6 andcytochrome f.
• Ferredoxintransfers electrons from PS I directly to NADP thus completing the light – driven
electron transport process.
• A final component of thylakoids necessary for photophosphorylation is a complex of proteins- F1
ATPase (Coupling factor- CF).
• ATP synthesis is stimulated by light-driven electron transport and electron transport in turn is
favoured by photophosphorylation.
• Thus CF couples the 2 processes together.

Photosystem I and II
▪ Step 1 – light excites e- in photosystem II
▪ Step 2 – e- move to primary e- acceptor
▪ Step 3 – e- move along electron transport chain (ETC)
▪ Step 4 – light excites e- in photosystem I
▪ Step 5 – e- move along 2nd (ETC)
▪ End – NADP+ combine H+ to make NADPH
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 ▪ P680 Pheo PQA PQB Cyt b6/f complex PC P 700
▪
P700 A0 (Chl) A1 (Quinone) FeSx FeSA FeSB Fd FNR NADP+
Chemiosmosis powers ATP synthesis in the light reactions
• The intra thylakoid energy of the elctrochemical potential gradient of protons (high H+ inside
thylakoid and low H+ in stroma) is the driving force behind the conversion of ADP and Pi into ATP,
a process catalyzed by an ATPase, located in the thylakoid membrane.
• ATP synthase – multifunctional protein - carrier protein that harnesses energy in thylakoid
membrane and it catalyzes ATP.
• This electron transport is responsible for generating most of the electrochemical proton gradient.

• ATP Synthase converts potential energy of protons (concentration gradient) into chemical energy of
ATP across thylakoid membrane. H+ move down their concentration gradient forming ATP
from ADP

• The electron transport chains are arranged with the photosystems in the thylakoid membranes and
pump H+ through that membrane

• The flow of H+ back through the membrane is harnessed by ATP synthase to make ATP
• In the stroma, the H+ ions combine with NADP+ to form NADPH
• This is known as the chemiosmotic theory based on proton motive force – proposed by Mitchell.
Cyclic photophosphorylation Non-cyclic photophosphorylation
1. In this process only PS I is functional 1. Both PS I and PS II are functional
2. Electron travels in a cyclic manner 2. Electron travels in a non- cyclic manner
3. Electron returns back to the chlorophyll 3. Electron is finally accepted by NADP+
4. Assimilatory power, ATP only is formed. 4. Assimilatory powers, ATP and NADPH are formed.
5. Oxygen is not evolved 5. Oxygen is evolved as a by-product
6. Photolysis of water is absent 6. Photolysis of water is present
7. It occurs at low light intensity, less 7. It performs best under optimum light intensity,
availability of CO2 and under anaerobic presence of CO2 and under aerobic conditions
conditions
8. Most of it occurs in stroma thylakoids 8. It occurs in grana thylakoids
9. It is not inhibited by DCMU 9. It is inhibited by DCMU as it blocks electron
(Dichlorophenyl dimethyl urea) transfer between PS II and PS I
10. The system is found dominant in 10. The system is found dominant in green plants.
photosynthetic bacteria.

Summary—Light Dependent Reactions

a. Overall input
light energy, H2O.
b. Overall output
ATP, NADPH, O2.
NADPH is the final (terminal) product of chloroplast light-reaction.
ATP

Princy Mol A P, Assistant Professor, Dept. of Botany 16