Photosynthesis in Higher Plants

1. Definition: “The physico-chemical

process in which sunlight drives the synthesis
of organic compounds is known as
photosynthesis”
Or
The Anabolic and Endergonic process in which
complex organic material i e Glucose is formed from
simple inorganic substance like CO and H O in the
2 2
presence of Sunlight with help of Chlorophyll in
which O is evolved as byproduct.
2

2. Features Of Photosynthesis in
higher plants (Oxygenic Photosynthesis):
i. It is physicochemical process
ii. Carbon dioxide and water are used as raw
material.
iii. Sunlight is trapped with the help of
chlorophyll pigment.
iv. Glucose is formed & Oxygen evolved as
byproduct.
v. It is uphill, anabolic, endergonic red-ox
process.
3. Importance of Photosynthesis:
i. Produce food for almost all organisms on the
earth except hydrothermal vent ecosystem
organisms.
ii. Produce oxygen & absorb carbon dioxide.
iii. It is Basis of the life on the earth.

4. What Do You Know & Early

Experimentation:
We have learn about photosynthesis in early classes
through some simple experiments shows that
chlorophyll (green pigment of the leaf), light and
CO are required for photosynthesis to occur.
2

1. Variegated leaf Experiment:

i. In this experiment we had Looked for starch
formation in a leaf. a variegated leaf or a leaf
that was partially covered with black paper and
partially exposed to light.
ii. On testing these leaves for starch it was clear
that photosynthesis occurred only in the green
parts of the leaves in the presence of light.

2. Moll’s half leaf Experiment:

 i. It is known as the half-leaf experiment, where a
part of a leaf is enclosed in a test tube (wide
mouth bottle) containing some KOH soaked
cotton (which absorbs CO ), while the other half
2
is exposed to air.
ii. The setup is then placed in light for some time.
On testing for starch later in the two halves of
the leaf, the exposed part of the leaf tested
positive for starch while the portion that was
in the tube, tested negative.
iii. This shows that CO is required for
2
photosynthesis.
3. Joseph Priestley's Experiment:

In 1770 he performed a series of experiments that

revealed the essential role of air in the growth of
green plants. He also discovered oxygen in 1774.
 • Priestley's observation :
i. A candle burning in a closed space i.e. a bell
jar, soon gets extinguished.
ii. Similarly, a mouse would soon suffocate in a
closed space.
• Priestley's conclusion :
He concluded that a burning candle or an
animal that breathe the air, both somehow,
damage the air. But when he placed a mint
plant in the same bell jar, he found that the
mouse stayed alive and candle continued to
burn.
• Priestley's hypothesis :
“Plants restore to the air (O2) whatever
breathing animals and burning candles
remove.”

4. Jan Ingenhousz's Experiment:

i. Using a similar setup as the one used by
Priestley but by placing it once in the dark and
once in the light, he showed that sunlight is
essential to the plant process (photosynthesis)
that somehow purifies the air fouled by burning
candles or breathing animals.
ii. In an another elegant experiment with an aquatic
plant showed that in bright sunlight small
bubbles were come out from the green parts,
while in the dark they did not.

In Light In Dark

• Conclusion :
It is only the green parts of the plants that could
release oxygen in the presence of sunlight.

5. Julius Von Sachs Experiment:

i. He provided evidences for production of glucose
when plants grow. Glucose is usually stored as
starch.
ii. His later studies showed that the green substance
in plants is located in special bodies within plant
 cells. (Today the green Colour substance is
known as chlorophyll and special bodies are
chloroplasts)

6. T W Engelmann’s Experiment:
i. He performed his experiment on Cladophora a
green algae & aerobic bacteria.
ii. He put the Cladophora in tiny glass tube in water
with aerobic & motile bacteria.
iii. He split the light using glass prism into seven
colours and exposed it to the Cladophora tube
setup.
 • Observation: The aerobic bacteria moves and
gathered in the regions of blue and red light.
• Conclusions: Blue and Red light is important
for photosynthesis.
• Note: This is First Action Spectra of
Photosynthesis.
7. Cornelius Van Neil Experiment:
He was microbiologist and perform his
experiment on purple and green bacteria.
• Conclusion:
i. Photosynthesis is essentially a light dependent
reaction in which hydrogen from a suitable
oxidisable compound reduces carbon di-oxide to
carbohydrates.
ii. Also Concluded that water is hydrogen donor in
green plants. Proved By Ruben & Kamen.
iii. Oxygen evolved from water & Not from CO2.

7. Van Neil’s Equation:

LIGHT
2H2A + CO2 2A + CH2O +H2O
• Current Equation of Photosynthesis:
LIGHT
6CO +12 H O C H O +6H O+6O
2 2 6 12 6 2 2
 5. Site of Photosynthesis:
i. Photosynthesis occurs in the green parts of the
plants like leaves, sepals, juvenile or herbaceous
stem, Cladode, phylloclade etc.
ii. Commonly photosynthesis occurs in the
mesophyll cells of the leaves.
iii. Mesophyll cells are chloroplast containing
parenchymatous cells present in leaves.
iv. Inside mesophyll cells chloroplast perform
photosynthesis.
v. v. Chloroplast is lens shaped green colored
double membrane bound cell organelle.
vi. vi. It contains the membranous structure
suspended in the stroma known as grana.
vii. vii. Grana is stack of coin like structure made of
many stacks of membranous sacs (Thylakoids).
viii. viii. Each Stack of thylakoid is known as
granum.
ix. ix. Granum are interconnected with stromal
lamellae.
x. x. Each granum consist of many thylakoids
arranged like stack.
 xi. xi. A thylakoid is membranous sac consist of
membrane and Lumen (Cavity).
xii. xii. The membrane of thylakoid is very
important as posses photosynthetic pigments
arranged in Photosystems, Cytochromes
(Electron Carrier) & ATP Synthases etc.
xiii. xiii. The colorless fluid surrounds the grana is
known as stroma.
xiv. xiv. The chloroplast shows clear division of
labor, the grana involved in trapping sunlight
and converting it into chemical energy i e ATP &
NADPH+H+ (Light Reaction) while the stroma
use these chemical energy to produce sugar
through enzymatic reactions (Dark Reaction).

Fig: Site of Photosynthesis

 ❖ Quantasome.
i. The unit of photosynthesis present in the
membrane of thylakoids known as Quantasome.
ii. It was discovered by Rodrick Park in1962.
iii. It consist of photosynthetic pigments and redox
carriers.
iv. There are two types of them the small and large
Quantasome.
v. After discovery of photosystems they rarely
used in scientific literature.
❖ Alignment of Chloroplast.
i. The chloroplast align themselves inside the
mesophyll cells to get optimum light whole the
day.
ii. The alignment of chloroplast changes with
change in light intensity.
iii. Early in the morning or late in the evening when
the light intensity is quite low chloroplast align
themselves perpendicular to the incident light to
get optimum light known as Epistrophe.
iv. In the Afternoon when the light intensity is very
high chloroplast align themselves parallel to the
incident light or to the transverse walls to avoid
Photoxidation known as Parastrophe.
v. In the moderate light condition chloroplast
shows random alignment known as Apostrophe.

Fig: Alignment of Chloroplast

Fig: Alignment of Chloroplast

Fig: Alignment of Chloroplast

Q. When do you think the chloroplasts
will be aligned with their flat surfaces
parallel to the walls?
Ans: In High light intensity
Q. When would they be perpendicular
to the incident light?
Ans: In Low light intensity
In Text Questions

6. Photosynthetic Pigments:
i. Pigments are substance that absorbs light of
specific wavelength and imparts specific Colour.
ii. The pigments involved in photosynthesis are
known as photosynthetic pigments.
iii. Different parts of plant shows different bright
colours due to pigments.
iv. Different pigments can be separated with the
help of paper chromatography.
v. The main photosynthetic pigments found in
plants are Chlorophylls & Carotenoids.
vi. Some bacteria possess Phycobillins along with
Bacteriochlorophyll and Bacterioviridin.
Let’s see one by one:
A. Chlorophylls:
i. The green colored photosynthetic pigment
present in the chloroplast are known as
chlorophylls.
ii. It is the most active & abundant pigment found
in all photosynthetic plants.
iii. It is lipid in nature and insoluble in water but
soluble in organic solvents.
iv. There are following 7 types of chlorophylls
found in photosynthetic organisms, i e
Chlorophyll-a, b, c, d, e, Bacteriochlorophyll and
Bacterioviridin.
a) Chlorophyll-a: (C H O N Mg)
55 72 5 4

i. It is the bluish green or bright green

pigment.
ii. It is found in all photosynthetic organisms
so known as universal pigment.
iii. It is also known as primary or essential
photosynthetic pigment as it act as reaction
 center in photosystem and can convert light
energy into chemical energy.
iv. It require sunlight and Mg, Fe & N for
synthesis from glycine & succinyl Co-A or
Glutamic acid.
v. Chemical structure made up of a Porphyrin
head & a Phytol tail.
vi. Porphyrin head is hydrophilic and absorb
light while Phytol tail is hydrophobic and
remain embedded in lipid bilayer of
thylakoid membrane.
vii. Imperial formulae is C H O N Mg.
55 72 5 4
viii. Mostly absorb blue violet & red light.
b) Chlorophyll-b (C H O N Mg)
55 70 6 4

i. It is yellow green pigment found in

photosynthetic plants.
ii. It is structurally similar with chlorophyll-a
except one -CH group in the place of –CHO.
3
iii. It absorbs blue-violet & red light.
B. Carotenoids:
i. They are yellow orange colour pigments
found in chloroplasts & chromoplasts.
ii. They are universal in occurrence (Except
Eubacteria) & insoluble in water.
 iii. They absorbs blue-violet light.
iv. They are of two main types i e Carotenes &
Xanthophylls.
v. Carotenes are yellow orange in color and
they are pure hydrocarbons C H .
40 56
vi. Most common carotenes are beta-carotenes.
vii. Xanthophylls are yellow colored
carotenoids.
viii. They are oxygenated hydrocarbons
C H O.
40 56 2
ix. The common xanthophyll is Lutein.
x. Lutein is responsible for autumn colour of
leaves.
xi. Diatoxanthin & Fucoxanthin are rare
Xanthophylls.
xii. Carotenoids are accessory pigments and
protect photosystem from nascent oxygen.
xiii. Helps in entomophily & zoochory.
xiv. Beta-carotene is precursor of vitamin-A.
C. Phycobillins:
i. They are photosynthetic pigments found in
Red algae & Cynobacteria.
ii. They are found in the form of
Phycoerythrins (Red) in red algae &
Phycocynins (Blue) in Cynobacteria.
Photosynthetic Colour Empirical Chemical Nature
Formula
Pigment

Chlorophyll-a Bluish C55H72O5N4Mg Lipids

Green

Chlorophyll-b Yellow C55H70O6N4Mg Lipids

Green

Carotene Orange / C H
40 56 Hydrocarbons
Red

Xanthophyll Yellow C H O
40 56 2 Hydrocarbons

Note: All are soluble in organic solvents & insoluble in cold/normal

water

• Main Pigments in Higher Plants are.

• Chlorophyll-a
• Chlorophyll-b
• Carotene
• Xanthophyll
 7. Nature Of Light:
i. Sunlight is essential input for photosynthesis so
need to understand.
ii. Light is dual in nature as it propagates in the
form of waves but interact with matter in the
form of particles.
iii. Particle of lights are regarded as photons
(Corpuscles).
iv. Photons are discrete packets of energy.
v. The amount of energy with a photon is known
as quantum.
vi. The amount of energy present in the photon is
inversely proportional to the wavelength of the
light.
vii. Shorter the wavelength higher the energy.
viii. Among the whole spectrum of the light only
light of the wavelengths between 400 nm to 700
nm is visible to human eyes so known as visible
light.
ix. The same wavelength of light i e from 400 nm to
700 nm is found to use in photosynthesis so
known as Photosyntheticaly Active Radiations
(PAR).
x. Only visible light is suitable to use for
photosynthesis as possess exact amount of
 energy suitable for photosynthesis without any
damage to photosynthetic meachineary.
xi. The light of very low wavelength (UV) possess
too much energy so can damage the
photosystems & the light of higher wavelength
(Infra-Red) possess too low energy to carry
photosynthesis.

Fig: Visible Spectrum & PAR

8. Absorption & Action Spectra:
i. Upto this point we knows that among the
electromagnetic spectrum only visible spectrum
of light is useful for photosynthesis & known as
PAR.
ii. The visible spectrum of light also consist of
seven different colors (VIBGYOR) representing
different wavelengths.
iii. Scientist was eager to know exactly which color
band among visible light is more useful in
photosynthesis.
iv. Early such attempt was made by T. W.
Engelmann, using Cladophora, prism & aerobic
bacteria. He concluded that blue & red light is
most useful light for photosynthesis (First
Action Spectrum).
v. Later scientist study it in detail & formulate
Absorption & Action Spectrum.
vi. Absorption Spectrum: The graphical
representation of absorption of different
wavelengths of light by various photosynthetic
pigments is known as absorption spectrum.
 400 nm 500 nm 600nm 700nm

Fig: Absorption Spectrum

❖ Features of The Absorption Spectrum:

i. Blue & Red light shows high absorption.
ii. Blue light is highest absorbed light.
iii. Chlorophyll-a shows two absorption peaks one
higher in blue light region (430 nm) & other
smaller in red light region (662 nm).
iv. Chlorophyll-b also shows two absorption peaks
one higher in blue light region (450 nm) & other
smaller in red light region (644 nm).
v. Carotenoids shows twin peaks in blue region
(450-460 nm)
vi. Action Spectrum: The graphical representation
of rate of photosynthesis plotted against
different wavelengths of light is known as action
spectrum of photosynthesis.

Fig: Action Spectrum of Photosynthesis

❖ Features of the Action Spectrum:

i. Blue & Red light shows high rate of
photosynthesis.
ii. Blue light shows highest rate of photosynthesis.
 ❖ Overlap of Absorption Spectrum of chlorophyll-a and
Action Spectrum of photosynthesis:

i. The absorption spectrum of chlorophyll-a shows

overlap with action spectrum of photosynthesis.
ii. This overlap proves that blue & red light not
only absorbed but also used in photosynthesis.
iii. But this overlap is not one to one because many
other accessory pigments like Chlorophyll-b,
Carotene, Xanthophylls also involved in
photosynthesis along with Chlorophyll-a.
iv. The involvement of other accessory pigments
widen the absorption spectrum & increases
photosynthetic efficiency.
v. Also the accessory pigments protect
Chlorophyll-a from phot-oxidative damage.
Q. Can you guess which the most abundant
plant pigment in the world is?
Ans: Chlorophyll-a
Q. can you determine the wavelength (colour
of light) at which chlorophyll a shows the
maximum absorption? Does it show another
absorption peak at any other wavelengths
too? If yes, which one?
Ans: Yes, at blue light (430 nm), yes it also shows
a small peak at red light region (662 nm).
Q. By looking at Figure 13.3c can you say
that there is a complete one-to-one overlap
between the absorption spectrum of
chlorophyll a and the action spectrum of
photosynthesis?
Ans: No, because other accessory pigments
also involved in photosynthesis like
carotenes, xanthophyll, chlorophyll-b etc.

In text Questions
9. The Flashing Experiments:
i. Robert Emerson & William Arnold performed
experiment with the use of light known as
flashing experiment.
ii. They use green algae chlorella and artificial light
for their experiment.
iii. They illuminate the algae with light for different
interval of time in short flashes.
iv. They also calculate photosynthetic efficiency in
terms of quantum yield for different
wavelengths of light or colours.
v. They found some interesting results.
vi. The Red Drop: The decline in quantum yield or
photosynthetic efficiency towards the far red
end (more than 680nm) of the spectrum is
known as red drop.
 vii. Or Decline in the rate of photosynthesis when
chloroplast is illuminated with far red light
(More 680 nm) is known as the red drop.
viii. The Emerson's Enhancement Effect:
The increase in quantum yield or photosynthetic
efficiency when both long (more than 680nm)
and short wavelengths (680 nm) of light were
given together is known as Emerson Effect or
Emerson enhancement Effect.
ix. The Emerson's enhancement effect point outs
the presence of two types of photosystems the
one works at red light while other far red light.

10. Photosystem:
i. Photosystems are structural & functional unit of
photosynthesis.
ii. They are located in the membrane of grana &
stromal lamellae.
iii. They are composed of photosynthetic pigments
embedded in protein complexes.
iv. Every photosystem is made up of a reaction
center & a light harvesting complex (LHC).
v. The reaction center of the photosystem possess a
pair of essential chlorophyll-a molecules (One as
per NCERT text book) which activates only after
absorb specific wavelength of light.
 vi. The Light Harvesting Complex (LHC) or
Antenna Complex is made up of 250- 400
molecules of accessory pigments like
chlorophyll-b, carotenes, Xanthophylls etc.
vii. The chlorophyll-a molecules at reaction center
can absorbs light as well as can concert it into
chemical energy so known as essential pigments.
viii. The other pigments present in the LHC absorb
the light & transfer the energy to reaction center
molecules through resonance as they are not able
to convert it into chemical energy.
ix. LHC increases the photosynthetic efficiency and
also protect reaction center from Photoxidation.

Fig: Structure of Photosystem

 ❖ Types of Photosystem (PS):
i. There are two types of photosystems on the
basis of their absorption specificity i e PS-I &
PS-II.
ii. The name of photosystems are given as per the
sequence of their discovery.
iii. The PS-I possess the chlorophyll-a molecules at
their reaction center which absorb light at 700
nm & activates, so known as P700.
iv. The PS-II possess the chlorophyll-a molecules at
their reaction center which absorb light at 680
nm & activates, so known as P680.

Fig: Types of Photosystem

 ❖ Distribution of Photosystem (PS)
in Grana & Stromal lamellae:
i. The non-appressed parts of the grana which are
exposed to the stroma (End Membrane & Margins)
& stromal lamellae possess PS-I in abundance, but
lacks NADP reductase so can carry out only
Cyclic Photophosphorylation.
ii. The appressed parts of the grana which are
unexposed to the stroma (intergranal thylakoids)
possess PS-II.
iii. The appressed regions also possess NADP
reductase so can carry out non-cyclic
photophosphorylation.

Fig. Distribution of PS in Grana & Stromal

lamellae
Fig. Distribution of PS in Grana & Stromal
lamellae
 ➢Mechanism of Photosynthesis:
i. Photosynthesis is complex and multistep
process.
ii. Photosynthesis occurs in two stages or phases
at two different places in the chloroplast.

A. Light Reaction
B. Dark Reaction

A. Light Reaction:
i. It is an oxidation reaction in which water get
oxidized.
ii. It is also known as photochemical phase or Hills
Reaction.
iii. It occurs at grana of chloroplast.
iv. It was studied in detail by Hills & Bendall.
 ❖ Hills & Bendall Experiment:
i. They perform an experiment on isolated
chloroplast of Stellaria media using artificial
oxidizing agents like ferricyanide in the absence
of CO2.
ii. They observed that when the isolated chloroplast
illuminated with light in the presence of
oxidizing agents and absence of CO2, oxygen get
evolved.
iii. It demonstrated that oxygen evolves from water
& not from CO2
iv. They also used artificial hydrogen acceptor like
Potassium ferricyanide, Benzoquinone,
chromate and DCPIP & natural hydrogen
acceptor like NADP etc.
v. All this hydrogen acceptor can be named as
Hills Reagents.
 ❖ Ruben & Kamen Experiments:

i. Later Ruben & Kamen proves that oxygen

evolves from water & Not CO using radio
2
18
isotope of oxygen O to radiolabel water.
ii. Ruben & Kamen found that when water
radiolabeled oxygen evolved shows
radioactivity.
 ❖ What exactly occurs in light reaction?
i. In light reaction solar energy is converted into
chemical energy in the form of ATP & NADPH +
+
H
ii. Light reaction requires photosynthetic pigments
to trap sunlight.
iii. Light reaction includes following processes
which occurs sequentially..
(i) Absorption of sunlight by pigments.
(ii) Excitation of chlorophyll-a molecule @ reaction
center.
(iii) Splitting of water
(iv) Release of oxygen
(v) Formation of high energy intermediates ATP &
+
NADPH+H
 iv. For easy understanding we will study then as
four key processes as follows
1)The electron transport
2)Photolysis (Splitting) of water
3)The Photophosphorylations
4)Chemiosmosis

❖ The Basics of Electron Transport:

i. What is ETS: The Electron transport system is a
series of electron carrier complexes that transfer
electrons from electron donors to electron
acceptor via redox reactions.
ii. Where it occurs: It occurs mostly on the grana or
membrane of thylakoids as its most of the
components are present in the membrane of
thylakoid.
iii. What are the components of ETS: It must need
three things to work i e the electron donor,
electron acceptor & electron carriers.
iv. Water acts as electron donor and NADP+ acts as
terminal electron acceptor here in ETS.
v. Plastoquinone, Cytochromes, Plastocynin,
Ferrodoxin, Ferrodoxin Reducing Agents (FRS),
Ferrodoxin NADP Reductase (FNR) etc. are
intermediate electron carriers.

Fig: The ETS

vi. Plastoquinone is an isoprenoid quinone molecule

found in the hydrophobic region of the thylakoid
membrane.
vii. Cytochromes are heam (Fe++) containing large
protein complexes.
viii. Plastocynin is a copper containing protein
complex.
ix. PQ and PC are mobile electron carriers of ETS.
x. How ETS Starts: ETS starts with photoexcitation of
reaction center of PS-II & PS-I i e P680 & P700
respectively.
xi. Absorption of light can change the redox potential
by bumping an electron from low energy orbit to
high energy orbit known as photoexcitation.
xii. The P680 absorbs sunlight of 680 nm and get
excited while P700 absorb sunlight of 700 nm and
get excited.
xiii. The excited P680 & P700 lose their high energy
electron to primary electron acceptors located
nearby.
xiv. How ETS Progress: The emitted high energy
electron then moves through the electron carriers
before it reach to NADP.
xv. The driving force responsible for the movement
of the electron is redox potential.
 xvi. What is redox potential: It is a measure of the
tendency of a chemical species to acquire or lose
electron to another compound and thereby get
reduced or oxidized respectively.
xvii. Redox potential measured in mili volts (mV).
xviii. On redox potential scale higher the redox
potential stronger the tendency to gain electron
(Oxidized state) while lower the redox potential
stronger the tendency to lose electron (Reduced
state).

Fig: Redox Potential Scale

xix. Thus electrons can move from compound with
low redox potential to compound with high redox
potential.
xx. All the components of ETS are arranged from
their low redox potential to high redox potential
so can transfer electron the next component.
xxi. When an electron carrier lose its electron to the
next it get oxidized the electron carrier get the
electron get reduced.
1. The Electron Transport:
i. The P680 of PS II absorb red light of 680nm
which excites the electrons of its reaction
center.
ii. The excited electrons jump out of its orbit.
iii. This electron then picked up by an electron
accepter (Primary Electron Acceptor) which
passes them to an electron transport system.

Fig: ETS on Redox Scale

iv. The electron transport system consist of

electron carriers and cytochromes.
v. This movement of electron from primary
electron acceptor to the PS I is downhill in
terms of redox potential.
vi. The electron are not used up as they passed on
& then reach to the pigments of PS I.
vii. Simultaneously the PS I gives out excited
electrons which then accepted by the acceptor
molecule of higher redox potential.
viii. These electrons moved downhill again to NADP+
ix. This whole scheme of transfer uphill to the
acceptor of electron starting from the PS II,
downhill to PS I, excitation of electrons, transfer
to another acceptor and finally downhill to
NADP+ is Known as the Z scheme due to its
characteristic shape on a redox scale.

Fig: The ETS

 2. Photophosphorylation
i. Phosphorylation is the process of synthesis of
ATP.
ii. There are three types of phosphorylations
observed in the living world, they are as follows.

Phosphorylation

I. Substrate Level II. Oxidative III. Photo-

Phosphorylation Phosphorylation Phosphorylation

iii. In substrate level phosphorylation ADP take

phosphate from phosphorylated substrate.
iv. In oxidative phosphorylation an oxidisable
substrate get oxidized and the energy released
is used for synthesis of ATP from ADP &
inorganic phosphate (iP) eg. respiration.
v. In photophosphorylation energy from sunlight is
used for the synthesis of ATP from ADP &
inorganic phosphate (iP).
vi. There are two types of photophosphorylation
i.e.
A. Cyclic Photophosphorylation
B. Non-cyclic Photophosphorylation
A. Cyclic Photophosphorylation

i. When only PS I is functional it results into cyclic

flow of electron known as cyclic
photophosphorylation.
ii. It possibly occurs in the stromal lamellae as it
lack NADP Reductase enzymes and PS-II but
possess PS-I.
iii. It also occurs when only light of wavelengths
beyond 680 nm (700nm) are available for
excitation.
iv. It leads to synthesis of only ATP.
v. The electron return back to PS-I after one cycle.

B. Non-Cyclic Photophosphorylation
i. When both the photosystems work together in
series, first PS II & then PS I, a process called
non-cyclic photophosphorylation occurs.
ii. It results in unidirectional flow of electrons from
water to NADP as two photosystems are
connected by an electron transport chain (ETS).
iii. Non-Cyclic Photophosphorylation occurs in the
appressed part of thylakoid membrane (Granal
Thylakoid).
 iv. It leads to synthesis of both ATP as well as
+
NADPH + H known as assimilatory powers.
v. When the sunlight of 680 nm falls on PS-II
excites a pair of chlorophyll molecules present
in its reaction center.
vi. The excited chlorophylls then emits a pair of
electrons.
vii. The electrons then accepted by the primary
electron acceptor (Pheophytin) present towards
the outer side of thylakoid membrane.
viii. The electrons then transferred to plastoquinone
(PQ).
ix. The Plastoquinone transfers it to cytochrome-b6-
f complex.
x. Cyt-b6-f complex transfer it to plastocynin (PC).
xi. The electrons then taken by oxidized PS-I who
looses its electron to FRS after excited by the
exposure of sunlight of 700 nm along with PS-
II.
xii. The FRS then give that electrons to Ferrodoxin.
xiii. Finally the electrons taken by NADP+ which
then reduced to NADPH with the help of NADP
reductase (FNR).
xiv. The electron moves uphill from PS II to primary
electron acceptor, then downhill from primary
 acceptor to PS I through ETS, again uphill from
PS I to its primary acceptor & finally downhill
from primary acceptor to NADP.
xv. All the electron carriers when arranged on
redox potential scale sequentially the give Zig-
Zag shape so the scheme is known as Z-Scheme.

Fig: Non-Cyclic Photophoshorylation

 3. Photolysis of Water
i. The PS II needs continual supply of electrons as
it loses electrons to NADP continuously.
ii. This can be done by splitting of water.
iii. The oxidized PS II act as strong oxidizing agent
and split water molecules into electrons,
protons & Oxygen.
iv. This splitting of water due to light is known as
photolysis of water.
v. The water splitting complex or oxygen evolving
complex (OEC) is associated with the PS II to the
inner side of thylakoid membrane.
++
vi. The water splitting complex needs Mn, Ca &
-
Cl ions to work.
 vii. So the proton and oxygen formed likely to be
released in the lumen.
viii. Oxygen can be diffused through membrane of
thylakoid as small & uncharged but protons
can’t.
ix. As photolysis of water progress protons start to
accumulate in the thylakoid lumen which leads
to building their high concentration inside the
thylakoid lumen.
x. DCMU (3-(3,4-dichlorophenyl)-1,1-dimethylurea)
inhibit flow of electron from PQ to Cyt-b6-f
complex so can be used to block PS II in
herbicides.
xi. Paraquat stop flow of electron by inhibiting
Ferrodoxin so useful to block PS I in herbicides.

4e-
Q. Where are the protons and O formed likely to
2
be released in the lumen? Or on the outer side of
the membrane?
Ans: Protons & Oxygen both released and
accumulated in the lumen of the thylakoids as
water splitting complex is present towards the
inner side of the thylakoid membrane. Protons
can’t pass through the membrane as they are
polar in nature, while the oxygen can diffuse
freely through the membrane as small and
nonpolar molecule. In Text Question
4. The Chemiosmotic Hypothesis:
Fig: Chemiosmosis

i. This hypothesis explains how actually ATP are

synthesized.
ii. It was put forth by Peter D. Mitchell.
iii. This hypothesis states that “ATP synthesis is
linked to development of a gradient of proton
across a membrane.”
iv. Due to various processes the concentration of
+
proton (H ) increases inside the lumen while it is
decreases outside the lumen i e in stroma.
v. This difference in concentration of proton
generate gradient across the thylakoid
membrane.
vi. This gradient is used in the synthesis of ATP by
ATPase.
vii. The following Processes are responsible for the
formation of the gradient.
A) Splitting Of Water in the Lumen.
B) Pumping Of Protons from Stroma to Lumen
by PQ. PQ is like a solar energy operated
proton pump.
C) Removal Of Protons from Stroma by NADP
Reductase.

Fig: Thylakoid

viii. This three things results into two things,

(i) Creation of proton gradient or Proton
Motive Force across the thylakoid
membrane.
 H
(ii) Causes measurable decrease in the P
in the lumen.
ix. The breakdown of this gradient due to
movement of proton across the thylakoid
membrane releases energy which can be used
for ATP synthesis by the ATP synthase.
x. The ATP synthase or ATPase enzyme consist of
two parts i e F0 & F1.
Fig. ATPase

Stroma

Lumen

xi. The F0 is embedded in the membrane and forms

a transmembrane channel that carries out
facilated diffusion of protons across the
thylakoid membrane
xii. The F1 protrudes on the outer surface of the
thylakoid membrane on the stromal side.
xiii. The breakdown of gradient releases energy
which cause conformational changes in the
ATPase enzyme that catalases the formation of
ATP.
 Special Points
i. In the mitochondria the gradient is formed along
inner membrane between intermembrane space
and matrix. (High H+ @ intermembrane space &
Low @ Matrix).
ii. In the aerobic bacteria it forms along plasma
membrane between periplasmic space &
cytoplasm. (High H+ @ Periplasm & Low @
Cytoplasm).

B. Light Independent/Biosynthetic Phase

i. The products of light dependent phase are

Oxygen, ATP & NADPH.
 ii. The Oxygen diffused out of the chloroplast.
iii. The ATP & NADPH are used in the process of
synthesis of food or sugars.
iv. This phase of synthesis of sugars from CO2 &
H2O with the use of ATP & NADPH is known as
biosynthetic phase.
v. It is reductive phase of photosynthesis.
vi. It occurs in the stroma.
vii. This process is not direct depend on light but
require the products of the light phase i e ATP &
NADPH.
viii. The first product formed when CO2 is taken into
a reaction or fixed was discovered by Melvin
Calvin with the help of radioisotope of carbon
14
C.
ix. The First product identified was a three carbon
organic acid i e 3-Phosphoglyceric Acid(3-
PGA)
x. Later we find out that there is another group of
plants Who's first fixation product is a four
carbon organic acid i e Oxyalo Acetic Acid
(OAA).
xi. So there are two main pathways of CO2 fixation
i e C3 Pathway & C4 Pathway.
 xii. CAM is third & rare type of pathway of CO2
fixation found.
xiii. In C3 pathway the first stable product is three
xiv. Carbon acid i e PGA while in C4 pathways it is
four carbon acid i e OAA.
xv. After the discovery of first product it is also
important to find primary acceptor of carbon.
xvi. Scientists first tried to find a two carbon
compound but finally discovered a five carbon
compound Ribulose-Bis-Phosphate (RuBP).
xvii. Calvin discovered the entire pathway of carbon
fixation now named as Calvin Cycle.
1) Calvin Cycle / Photochemical Carbon
Reduction (PCR) Cycle / C3 Cycle:
i. Melvin Calvin & coworkers elucidate the detailed
pathway of synthesis of sugar.
ii. They perform experiment on chlorella using
14
radioisotope of carbon C.
iii. They find all the intermediate of the pathway
include first stable product 3-PGA & Primary
acceptor of carbon RuBP.
iv. They Get Nobel for their contribution.
v. They find that this process operate in cycle &
named as C cycle.
3
 vi. Later renamed as Calvin Cycle in the honor of
Melvin Calvin.
vii. Calvin Cycle occurs in all the photosynthetic
plants include C4 Plants.
viii. It can be described in three steps.
1. Carboxylation
2. Reduction
3. Regeneration
1. Carboxylation

i. In this step carbon dioxide is fixed into a stable

organic intermediate.
ii. RuBP accept CO and form two molecules of 3-
2
PGA.
iii. This reaction is catalyzed by RuBisCO.
iv. RuBisCO is an enzyme with dual activity.
2. Reduction
i. These are the series of reactions that lead to the
formation of glucose.
ii. The steps involves utilization of 2 molecules of
ATP for phosphorylation & 2 molecules of
NADPH for reduction per CO molecule fixed / 12
2
ATP & 12 NADPH per glucose molecule
produced.
iii. Produced glucose can be converted into sucrose
for transport & starch for storage.
iv. Also regarded as glycolytic reversal.
Notes
3. Regeneration

i. Regeneration of the primary carbon acceptor

molecule RuBP is essential for the continuity of
the cycle.
ii. Regeneration steps require one molecule of
ATP/RuBP.
NOTE:
 ➢ Energetics Of Calvin Cycle
i. The fixation of six CO and six turns of the cycle
2
requires for the synthesis of a glucose molecule.
ii. For fixation of one CO molecule 3 molecules of
2
ATP & 2 molecules of NADPH are required.
iii. It is probably to meet this difference in number
of ATP & NADPH used in the dark reaction that
the cyclic phosphorylation takes place.

Q. Work out how many ATP and NADPH

molecules will be required to make one
molecule of glucose through the Calvin
pathway.
Ans: Formation of one glucose molecule
requires six turns of Calvin Cycles, each turn of
cycle requires 3 ATPs (2 for phosphorylation & 1
for regeneration) & 2 NADPH+H+.
So 6 Turns x 3 ATP & 2 NADPH+H+ = 18 ATPs & 12 NADPH+H+.
Energetics of Calvin Cycle
In Out

Six CO One Glucose

2

18 ATP 18 ADP

12 NADPH 12 NADP
 Q. Work out how many ATP and NADPH
molecules will be required to make one molecule
of glucose through the Calvin pathway.
Ans: Formation of one glucose molecule
requires six turns of Calvin Cycles, each turn of
cycle requires 3 ATPs (2 for phosphorylation & 1
+
for regeneration) & 2 NADPH+H .
+
So 6 Turns x 3 ATP & 2 NADPH+H = 18 ATPs & 12
+
NADPH+H . In Text Question

❖ Photorespiration

i. RuBisCO is the most abundant enzyme in the

world.
ii. Its active site can bind to both CO & O .
2 2
iii. RuBisCO has a much greater affinity for CO when
2
the CO :O is nearly equal.
2 2
iv. Binding of RuBisCO for O2 or CO2 is
competitive & depend on their relative
concentration.
 v. In certain conditions RuBisCO bind to O2 &
perform photorespiration like High temperature,
High Light intensity, Low CO2 concentration etc
vi. A respiratory process in many higher plants by
which they take up oxygen in the light and give
out some carbon dioxide, contrary to the general
pattern of photosynthesis known as
Photorespiration.
vii. It has following names
• PCO (Photosynthetic Carbon Oxidation) Cycle
• C Cycle
2
• Glycolate metabolism
viii. In photorespiration RuBisCO release already
fixed CO2 rather than fixing it through
photosynthesis.
ix. Photorespiration starts in chloroplast but also
carried out in peroxisomes & mitochondria.
x. Photorespiration not result in synthesis of ATP
Or NADPH instead release already fixed CO2
with using ATP so a wasteful process.
xi. The biological function of photorespiration is
not known yet.
 • Warburg effect..
Decrease in the rate of photosynthesis by High O
2
Concentration known as Warburg effect.

Q. How Plant Avoid Photorespiration..

❖ Kranz Anatomy:

i. C4 Plants show Kranz Anatomy.

ii. In C4 leaves the mesophyll is not differentiated
into palisade & spongy tissue.it is homogeneous.
iii. In this leaves each vascular bundle is surrounded
by a ring or wreath of radially arranged large
bundle sheath cells.
 iv. Cells of mesophyll and bundle sheath are
interconnected by Plasmodesmata.
v. Bundle sheath cells lack intracellular spaces,
posses thick cell walls imperative for gaseous
exchange & having large number of chloroplasts.
vi. Mesophyll cells contains agranal chloroplast i e
chloroplast without grana. they are bigger in
size & more in number with abundant stroma.
vii. The chloroplast in other mesophyll cells contain
grana i e Granal chloroplast. they are small in
size with abundant grana and less stroma.
viii. Thus chloroplast are dimorphic in C4 plants.
ix. The enzyme PEPcase is present in mesophyll
cells while RuBisCO in bundle sheath cells.
 2) C4 Plants & C4 Pathway

i. These plants have the oxyaloacetic acid (OAA) as

the first CO2 fixation product.
ii. They are adopted to dry tropical regions.
iii. They use C3 pathway or Calvin cycle as main
biosynthetic pathway.
iv. The characteristics of C4 plants are as follows.
1. Special leaf anatomy.
2. Tolerance to high temperature.
3. Response to high temperature.
4. Lack photorespiration so more productive then
C3.
Eg. Sugarcane, Maize, Jowar, Amaranthus, Salsola,
Atripex etc.
 2) C4 Pathway
i. The primary CO2 accepter is 3 carbon molecule
Phosphoenol Pyruvate (PEP) present in
mesophyll cells.
ii. The enzyme responsible is PEPcase.
iii. Mesophyll cells of C4 lacks RuBisCO.
iv. The first stable product is C4 Acid OAA.
v. The C4 acid then converted into Malic Acid or
Aspartic Acid which then exported to bundle
sheath cells.
vi. In bundle sheath cells these C4 acid (Malic Acid)
is decarboxylated into a C3 acid(Pyruvate).
vii. The CO2 released then used in Calvin cycle by
RuBisCO.
viii. The C3 acid formed in decarboxylation
transported back to the mesophyll cells where it
phosphorylated back into PEP again.
ix. The bundle sheath cells are rich in RuBisCO but
lacks PEPcase.
x. Calvin Cycle is common in C4 and C3 plants.
xi. C4 pathway is also known as Hatch & Slack Or HSK
Pathway.
xii. C4 plants require 30 ATPs for synthesis of one
glucose molecule instead 18 ATPs in C3 Plants.
 3) CAM Plants & CAM Pathway
i. In some plants which grow in xerophytic
conditions the stomata remain closed during day
to check water loss due to transpiration this
plants shows different photosynthesis
mechanisms known as CAM.
ii. Such plants are known as CAM plants.
iii. Eg Xerophytic Succulents.(Crasusulaceae family)
like Kalanchoe, Bryophyllum, Opuntia, Agave,
Aloe, Euphorbia, Pineapple, Welwitschia.
iv. In these plants CO2 is taken during night when
stomata are open by PEP and OAA is formed in
the presence of PEPcase.
v. OAA then get reduced to malic acid thus malic
acid accumulated during the nights.
vi. During day when stomata are closed malic acid
get decarboxylated & release CO2 which then
used in Calvin Cycle to produce sugar.
vii. The pyruvate also get converted in starch thus
starch get accumulated during the day.
viii. Thus CAM plants perform C3 during day and C4
during night.
ix. The stomata open during night but remained
closed during day is known as Scotoactive
Stomata.
x. CAM plants also require 30 ATP & 12 NADPH
for synthesis for one glucose molecule.
 ❖ Factors Affecting Rate of
Photosynthesis
i. The rate of photosynthesis is very important in
determining the yield of plants including crop
plants.
ii. There are several factors which determine the
rate of photosynthesis.
iii. These factors can be divided into two types..
• A. Internal Factors
• B. External Factors
A. Internal Factors:
i. The factors which are related to plants are
known as internal factors.
ii. This factors determined by plants growth status
& genetic constitution.
iii. This factors includes following.
a. Number of Leaves.
b. Size of Leaves.
c. Orientation of Leaves.
d. Mesophyll Cells.
e. Number of Chloroplast.
f. Amount of Chlorophyll.
g. Age of Leaves.
 h. Internal CO2 Concentration.
B. External Factors:
i. The factors which are related to external
environment are known as external factors.
ii. This factors are climatically determined.
iii. This factors includes following.
a. Light
b. CO2 Concentration
c. Temperature
d. Water

• Blackmans Law of Limiting Factors

“If a chemical process affected by more
than one factor then its rate will be
determined by the factor which is nearest
to its minimal value.it is the factor which
directly affects the process if its quantity is
changed.”
 1. Light:
i. Light is basic requirement of photosynthesis.
ii. Light affects photosynthesis in three ways
a. Light Quality
b. Light Intensity
c. Duration of Exposure
a. Light Quality:
i. Spectral quality of light is important for
photosynthesis.
ii. Light of only 400-700 nm is useful for
photosynthesis & known as PAR.
iii. PAR consist of light of seven different
colours representing different wavelengths i
e VIBGYOR.
iv. Most effective spectral components are
blue & red while least effective is green.

b. Light Intensity:
i. Light intensity is also important for
photosynthesis.
ii. There is liner relationship between light
intensity & CO2 fixation rate at low light
 intensities because in this condition light
becomes limiting factor.
iii. At higher light intensity the CO2 fixation rate
doesn't affected by light intensities as other
factors become limiting.
iv. Light Saturation Point: The light intensity
beyond which no further increase occurred in
rate of photosynthesis.
v. Rate of photosynthesis saturates at 10 % of the
full sunlight.
vi. Very high intensities of light may cause
solarization to photosynthetic meachineary.
vii. Solarization includes phot-oxidative damage to
pigments, enzymes & photosynthetic carbon
compounds like RuBP.
viii. Light Compensation Point: The intensity of light
at which rate of photosynthesis equals to rate
of respiration in plants.
ix. At light compensation point there is no need of
gaseous exchange with atmosphere.
 c. Duration of exposure:
i. Duration of light exposure never affects rate of
photosynthesis if intensity of light & quality
remains constant.
ii. Intermitting light shows high rate of
photosynthesis as compare to continuous light
as ATP & NADPH2 get accumulated during
continuous light and its production get feedback
inhibited.
iii. Light period more than 12 hours even doesn’t
affect photosynthetic productivity.
 2. Carbon dioxide concentration
i. Carbon-di-oxide is the major limiting
factor for photosynthesis. The
concentration of CO2 is very low in the
atmosphere i.e., 0.03 to 0.04 percent or
300 to 400 PPM.
ii. C4 plants show saturation at about 360
–1
ul L while C3 responds to increased CO2
concentration and saturation is seen
–1
only beyond 450 ul L . Thus, current
availability of CO2 level (0.03 to 0.04%) is
limiting to the C3 plants
iii. Increased in CO2 concentration upto
0.05% can cause an increase in CO2
fixation rates but beyond this the levels
can become damage in over longer
periods.
iv. Green house crops/C3 plants such as
tomatoes and bell pepper show higher
photosynthetic rates under CO2 enriched
 atmosphere that leads to higher yield.
This phenomenon is called CO2 fertilizing
effect.
v. CO2 compensation point : It is the point
where rate of photosynthesis become
equal to the rate of respiration (NPP is
zero).
vi. CO2 compensation point for C4 plants is
0-10 ppm.
vii. CO2 compensation point for C3 plants is
25-100 ppm.
 3. Temperature
i. The dark reaction being enzymatic, is
temperature controlled as enzymes are
temperature sensitive.
ii. Though the light reaction is also temperature
sensitive but they are affected to a much lesser
extent.
iii. The C4 plants respond to higher temperature
(30 – 40°C) and show higher rate of
photosynthesis while C3 plants have a much
lower temperature optima (20 – 25°C).
Optimum temperatures also depends on the
habitat. Tropical plants have a higher
temperature optimum than the plants adapted
to temperate climates.

4. Water
i. Water is one of the reactant of the
photosynthesis (Light Reaction).
ii. Water affect photosynthesis indirectly through
its effect on the plants.
iii. Water stress can cause stomatal closure which
results in reduction in CO2 availability.
iv. Water stress also causes wilting of leaves which
reduces surface area available for
photosynthesis and hence decreases
photosynthesis.
➢ Numerical Basics
Notes