CHAPTER 13

PHOTOSYNTHESIS
IN
HIGHER PLANTS
 Introduction
• "Photosynthesis is a physico-chemical or photo-biochemical
process in which the organic compounds (carbohydrates) are
synthesized from the inorganic raw material (H2 O & CO2 ) in the

presence of light & pigments.

• O2 is evolved as by product or one of the net products".

• Light energy is converted into chemical energy by

photosynthesis.
 Introduction
• Photosynthesis is an :
• Anabolic (synthesizing) &
• Endergonic (Energy absorbing) process.
• Photosynthesis is a redox reaction during which oxidation of
H2 O occurs (as it provides H+ and e.) during light reaction and

reduction of CO2 occurs (as it accepts H+ & e.) during dark

reaction (biosynthetic phase).

Reduction of CO2

CO2 + H2O C6H12O6 + O2

Oxidation of H2O
 What Do We Know ?
• Look for starch formation in two leaves – a variegated leaf or a
leaf that was partially covered with black paper, and exposed to
light.
• On testing these leaves for the presence of starch, it was clear
that photosynthesis occurred only in the green parts of the
leaves in the presence of light.
 • .
Moll’s Half Leaf Experiment

• Conclusion:
• This showed that CO2 was required for photosynthesis.
Early Experiments (Joseph Pristley)
• Joseph Priestley (1733-1804) in 1770 performed a series of
experiments that revealed the essential role of air in the growth
of green plants.
• Priestley hypothesized as follows: “Plants restore to the air
whatever breathing animals and burning candles remove”.
• Priestley, discovered oxygen in 1774.
Early Experiments (Jan Ingenhousz)
• Using a similar setup as the one used by Priestley, but by
placing it once in the dark and once in the sunlight.
• Jan Ingenhousz (1730-1799) showed that sunlight is essential
to the plant process that somehow purifies the air fouled by
burning candles or breathing animals.
Presence Absence
of light of light
Early Experiments (Jan Ingenhousz)
• Ingenhousz in an elegant experiment with an aquatic plant
showed that in bright sunlight, small bubbles were formed
around the green parts while in the dark they did not.
• Later, he identified these bubbles to be of oxygen. Hence he
showed that it is only the green part of the plants that could
release oxygen.
Early Experiments (Julius Von Sachs)
• Julius von Sachs (1854) provided evidence for production of
glucose when plants grow.
• His later studies showed that the green substance in plants
(chlorophyll as we know it now) is located in special bodies
(later called chloroplasts) within plant cells.
• He found that glucose is made in green parts of plants and it is
usually stored as starch.
 Early Experiments
Early Experiments (Julius Von Sachs)
Early Experiments (Cornelius Van Neil)
• Cornelius van Niel (1897-1985): - A microbiologist
• Based on his studies on purple and green bacteria (milestone
contribution), he demonstrated that photosynthesis is essentially a
light-dependent reaction in which hydrogen from a suitable
oxidisable compound reduces carbon dioxide to carbohydrates.
• In green plants, H2O is the hydrogen donor and is oxidised to O2.
• Some organisms do not release O2 during photosynthesis. When
H2 S, instead is the hydrogen donor for purple and green sulphur
bacteria, the ‘oxidation’ product is sulphur or sulphate depending on
the organism andLight
not O2 .
CO2 + H2S CH2O + S

• Hence, he inferred that the O2 evolved by the green plant comes

from H2 O, not from carbon dioxide.
CO2 + H20 Light
CH2O + O2
 Early Experiments (Robert Hill)
• Detailed study of light reaction in
isolated chloroplast of Stellaria
plant. He illuminated the isolated
chloroplasts of Stellaria media in
the presence of hydrogen acceptors
(ferricyanides) in the absence of
carbon dioxide.
• The chloroplasts evolved oxygen

Robert Hill & Bendall

• They are credited for detailed study of light reaction and proposed
Z scheme.
Early Experiments (Ruben & Kamen, 1941)
• Then used O18 (Radioisotope technique) to show experimentally
that O2 in photosynthesis release from water.
Light
6CO2 + 12H2O18 C6H12O6 + 6H2O + 6O218
Where Does Photosynthesis Take Place ?
• Photosynthesis does take place in green leaves of plants, but it does so also
in other green parts of plant. Outer membrane
Inner membrane

Stroma lamellae

Granum

Stroma
Ribosome

Lipid droplets
Starch granules
Division of Labour In Chloroplast
Division of Labour In Chloroplast
Division of Labor
Alignment of Chloroplasts
• Usually the chloroplasts align themselves along the walls of
the mesophyll cells, such that they get the optimum quantity
of the incident light.
Conditions Alignment of chloroplasts
Parallel to the incident light / Lateral
High light intensity
walls (Parastrophe)
Perpendicular to the incident light
Low light intensity
(Epistrophe)
Random (Apostrophe)
Moderate light intensity
How Many Pigments Are Involved In Photosynthesis
Photosynthetic Pigments are of following types :
1. Chlorophyll 2. Carotenoids 3. Phycobillins
1. Chlorophylls :-
• Green Colored Pigment
• Light is required for their synthesis
• Soluble in organic solvents
Chlorophylls are of following types :
i. Chl-a : Universal Pigment (Found in all O2 liberating photosynthetic
Organisms)
Blue Green Colour in Chromatogram
ii. Chl-b : Accessory photosynthetic pigment found in Euglenoids,
Green Algae and Higher plants.
Yellowish green in chromatogram.
iii. Chl-c
iv. Chl-d
v. Chl-e
 Photosynthetic Pigments
 Chlorophyll-a C55H72O5N4Mg CH3 Group in IInd pyrrol ring

 Chlorophyll-b C55H70O6N4Mg CHO Group in IInd pyrrol ring

Chlorophyll (Tadpole like)
Structure of Chlorophyll :-

Porphyrine Head Phytol Tail (C20H39OH)

• Size = 15 x 15 Å • Size = 20 Å
• It is Hydrophilic in nature • It is Hydrophobic
• It consist of four N- • Tail remains
containing pyrrol ring embedded in lipid
(together called as tetra bilayer of thylakoid
pyrrol ring). membrane.
• All the N of pyrrol ring
 Phytol tail is absent
connected with central in Chl-c.
structure with Mg present in
the Chlorophyll
centre. synthesis :
Succinyl CoA + Glycine Protochlorophyll Light
Chlorophyll
2H
• This reaction is catalyzed by Iron (Fe)
Photosynthetic Pigments
Functions of carotenoids
1) They are accessory pigments and make photosynthesis more
efficient by absorbing different wavelengths of light.
2) They protect chl-a from photo oxidation and they also protect
photosynthetic machinery by converting lethal nascent oxygen
into unharmful molecular oxygen, Thus, also called shield
pigments.

(3) β-carotene is acts as a precursor of vitamin-A

(4) They help in entomophily and zoochory.

 Phycobillins
3. Phycobillins :
• They are hot water soluble pigment
• They lack Mg and phytol tail.
Types of phycobillins:
i. Phycocyanin (Blue)
ii. Phycoerithrin (Red)
iii. Allophycocyanin (Light blue)
• They occur exclusively in BGA and Red algae as an accessory
pigments.
 Absorption Spectrum
• Graph showing the ability of pigments to absorb lights of
different wavelengths (absorption spectrum).
• Chl a shows maximum absorption at blue light. It show another
absorption peak at red light.
 Action Spectrum
• The graphic curve depicting the relative rates of photosynthesis
at different wavelengths of light is called action spectrum.
Figure showing the wavelengths at which photosynthesis occurs
in a plant (Action spectrum) .
Absorption Spectrum Of Chl-a Superimposed With Action Spectrum Of Photosynthesis
• Figure show that the wavelengths at which there is maximum
absorption by chlorophyll a, i.e., in the blue and the red regions,
also show higher rate of photosynthesis.
• Hence, we can conclude that chlorophyll a is the chief pigment
associated with photosynthesis.

Action

Absorption
Conclusion of Graphical Studies
• These graphs a, b and c together show that most of the
photosynthesis takes place in the blue and red regions of the
spectrum; some photosynthesis does take place at the other
wavelengths of the visible spectrum.
• It happens because other thylakoid pigments like chlorophyll b,
xanthophylls and carotenoids, which are called accessory
pigments, also absorb light and transfer the energy to
chlorophyll a.
• Indeed, they not only enable a wider range of wavelength of
incoming light to be utilised for photosynthesis but also protect
chlorophyll a from photo-oxidation.
T. W. Engelmann’s Experiment
He proposed first action spectrum of photosynthesis.

• It resembles roughly
the absorption spectra
of chlorophyll a and b.
Mechanism of Photosynthesis
There are two types of reactions in photosynthesis .
1. Light reaction (Takes place in grana & stroma lamellae)
2. Dark reaction ( Takes place in stroma)

What is Light Reaction

Light reaction or photochemical process includes:
1. Light absorbtion
2. Water splitting
3. Oxygen release
4. Synthesis of high energy chemical intrermediates like ATP & NADPH
Red Drop Effect and Enhancement Effect
• Red drop effect:
• Emerson and Arnold, while working on Chlorella by providing
monochromatic light (wavelength of single wavelength)
observed minimum photosynthetic yield, when supplied
monochromatic beam of 700 nm or > 680 nm only and called it
red drop.
• Enhancement effect or Emerson effect :
• They also observed enhancement in photosynthetic yield when
a mixture of light of wavelength i.e. both 700 nm and 680 nm
supplied together and called it enhancement effect.
Conclusion :
• Two types of photosystems (PS-I and PS-II) exist in
photosynthetic units. They operate simultaneously and their
operation / activation required 700 nm and 680 nm radiation.
 Photosystems
Quantasome (250 – 400 pigment molecule)

Lumen

Primary acceptor
Thylakoid F0
F1 Thylakoid
Membrane
e-
Photon Reaction
centre
Pigment Light harvesting complex
Molecules (LHC)
 Photosystems
On the basis of Reaction Centre, there are two types of Photosystem :
1. Photosystem – I (PSI)
• Reaction Centre = P700

2. Photosystem – II (PS II)

• Reaction Centre = P680
• These are named in the sequence of their discovery, and not in the
sequence in which they function during the light reaction
Light ( 680nm or below)
Both PSI and PSII Active Light ( More than 680nm)
Only PS I active
Distribution of Photosystems
• The PS II is located in the appressed region of granal thylakoids
and PS I in non appressed region of grana and in stroma
thylakoids. (In the other way we can say that granal thylakoids
have both PS I and PS II whereas stroma thylakoids have only
PS I).
Photophosphorylation
Synthesis of ATP from ADP and inorganic phosphate (iP) with the
help of light energy is known as photophosphorylation. It is of two
types :-
1. Non-cyclic photophosphorylation
2. Cyclic photophosphorylation

1. Non-cyclic photophosphorylation
• Both PS-I and PS-II are involved
• It occur on the membrane of grana thylakoids
• In this process:
• Photolysis of water takes place
• Oxygen release occurs
• ATP & NADPH synthesis take place
 Non- cyclic Photophosphorylation
LIGHT
H STROMA
Cyt
PQH b6 e
e PSI e-
-

e &f
PQ -

e -
H PC NADP+ H
LIGHT
HYTE
N Fe-Se Fd FNR
OP
PHE E NADPH + H
-

e - HO PROTON GRADIENT
H
Photolysis
OEC H ADP + iP
PS II ½ H PHOTO-
O PHOSPHORYLATION
e F F
ATP
-

Fd= Ferredoxin
LUMEN LI PID FNR= Ferredoxin NADP Reductase
H O R
P
THYLAKOID MEMBRANE H OSILAYE PQ= Plastoquinone
P B PC= Plastocyanin
OEC= Oxygen evolving complex
 Chemiosmotic Hypothesis
• Proposed by Peter Mitchell and were awarded by Nobel Prize.
• He proposed the mechanism of ATP synthesis in both chloroplast and
mitochondria.
• According to him, ATP synthesis is linked to development of proton
gradient (difference of H+ ion concentration) across membrane.
• Chemiosmosis require:
• A membrane
• A proton pump
• Proton gradient
• ATP synthatase enzyme complex (F0-F1 particle)

• The step that causes proton gradient to develop –

• Photolysis of water
• Proton pumping by plastoquinone (PQ)
• Reduction of NADP+ to NADPH in stroma
Chemiosmotic Hypothesis

(Ferredoxin NADP Reductase)

 Z-scheme
Low High

Redox potential Energy

High Low

• This whole scheme of transfer of electrons, starting from the PS II,

uphill to the acceptor, down the electron transport chain to PS I,
excitation of electrons, transfer to another accepter, and finally down
hill to NADP+ is called the Z scheme, due to its characteristic shape.
• This Z shape is formed when all the carriers are placed in a sequence
on a redox potential scale.
Quantum Requirement and Quantum Yield
Light (4 Photons)
Light 4e- 4e-
(4 Photons)
NADP +
4e-
PS
I 4e-
Thylakoid
Lumen Membrane
PS 4e- 4e-
II
OEC
2H20 4H+
Stroma
O2

Quantum requirement
• The number of light Quanta or photons required for the evolution of 1 mol. of O2 in
photosynthesis.
• Emerson calculated that the quantum requirement is 8.
Quantum Yield
• The number of oxygen molecule evolved by one quantum of light in photosynthesis
is called as Quantum yield.. Hence the quantum yield is 0.125 or 12.5%
Cyclic Photophosphorylation
• In cyclic photophosphorylation, only PS-I works.
• A possible location where this could be happening is in the
stroma lamellae which lacks PS II and NADP reductase enzyme.
• In cyclic ETS, no oxygen evolution occurs because photolysis of
water is absent.
• NADPH + H+ (Reducing power) is not formed in cyclic process.
There is formation of only ATP.
 Cyclic Photophosphorylation
Light
H+

e-
Cyt b6 e-
PQ
e- e- e- PS I
PQH2 Cyt f e-
PC Fd
H+ FeS
H+ e-
PROTON GRADIENT ADP + iP

F0
F1
THYLAKOID MEMBRANE
ATP
H+
Where are the ATP And NADPH Used ? (Dark Reaction/Biosynthetic Phase)

(i) In this process CO2 is reduced to sugar.

(ii) This process does not directly depend on the presence of light but is
dependent on the products of the light reaction like ATP and
NADPH2.
(iii) It is known as dark reaction but it doesn't mean that it occurs in dark
as it operates simultaneously with light reaction in presence of light.
(iv) Suppose, if light become unavailable, the biosynthetic process
continues for some time and then stops and after a while, if light
becomes available, then synthesis starts again.
Where are the ATP And NADPH Used ? (Dark Reaction/Biosynthetic Phase)

• Calvin studied the dark reaction in green algae Chlorella &

Scenedesmus. During his experiment he used radioisotopy (14C
radioisotope) and chromatography techniques for identification and
separation of intermediates of C3 –cycle
• The use of radioactive 14C by him in algal photosynthesis studies led
to the discovery that the first CO2 fixation product was a 3-carbon
organic acid i.e. Phosphoglyceric acid (PGA).
Primary Acceptor of CO2
Types of Dark reaction/ Biosynthetic phase
• C3 Plants = Calvin Cycle/ C3 Cycle
RuBisCO
• Primary CO2 acceptor = RUBP (5C)
Chloroplast
• First stable compound = PGA (3C)
Mesophyll Cell

Pepcase
2. C4 Plants = Hatch & Slack Pathway RuBisCO
• Primary CO2 acceptor = PEP (3C) Chloroplast Chloroplast
• First stable compound = OAA (4C) Mesophyll Bundle
Cell sheath cell
3. CAM Plants = CAM Pathway
• Primary CO2 acceptor = PEP (3C) RuBisCO Pepcase
• First stable compound = OAA (4C)
Chloroplast
Mesophyll Cell
Calvin Cycle or C3- Pathway
• Calvin & his co-workers then worked out the whole pathway and
showed that the pathway operated in cyclic manner.
• This pathway occurs in all photosynthetic plants; whether they
have C3 or C4 or CAM pathways.
• Calvin cycle can be described under three stages:
1. Carboxylation (Most crucial step)
2. Reduction
3. Regeneration
1. Carboxylation:
It is process of fixation of CO2 into stable organic intermediate
which is catalyzed enzyme RuBiSCO (Ribulose 1,5 bis phosphate
carboxylase oxygenase).
 Atmosphere

Ribulose 1,5 CO2 + H2O

bis phosphate RuBisCO
(Most Crucial
RuBP (5C) Carboxylation
step)
ADP
Keto Acid (6C)
Unstable

Regeneration 2x3 Phosphoglycerate

PGA (3C)

ATP 2ATP +
Reduction 2NADPH+ H +
2XPGAL (3C)
Triose
Phosphate
2ADP + 2NADP
Sucrose, Starch
C4 Pathway/Hatch & Slack Pathway
C4 pathway also called as:
 CO2 Concentrating Mechanism
 Co-operative Photosynthesis
 Dicarboxylic Acid Cycle (DCA Cycle)
• Most of the C4 plants are monocots (Tropical grasses), which
belong to Gramineae & Cyperaceae families. C4 plants are adapted
to hot and dry environment.
Eg. : C4 plants . Sugarcane, Maize, Sorghum.

• Wheat, Rice and Barley are C3 species.

C4 plants are special because :

(i) They lack a process called photorespiration so have greater

productivity of biomass.
(ii) They tolerate higher temperature
(iii) They show a response to high light intensities
(iv) They have a special type of leaf anatomy (Kranz anatomy)
 Kranz Anatomy
Kranz (Wreath) Anatomy- Present in leaves of C4 plants..
• The bundle sheath cells may form several layers around the
vascular bundles, they are characterised by –
i. Having a large number of chloroplasts
ii. thick walls impervious to gaseous exchange
iii. no intercellular spaces
 MESOPHYLL CELL BUNDLE SHEATH CELL

Epidermal cell
AGRANAL CHLOROPLAST

OAA MALIC MALIC

(4C) NADPH2
ACID (4C) ACID (4C)
NADP
NADP
Carboxylation/Fixation
PEPCase CO2
GRANAL CHLOROPLAST
Carboxylation/ NADPH2 CALVIN
Fixation CYCLE

2ATP
PEP (3C) PYRUVIC PYRUVIC
TRIOSE
Regeneration ACID (3C) ACID (3C)
PHOSPHATE

HCO3 - STARCH

SUCROSE
SUCROSE
CO2 PLASMODESMATA
 C4 Pathway
• C4 plants total 30 ATP and 12 NADPH2 are utilized for synthesis of
one glucose.
• In C4 plants photorespiration does not occur.
• This is because they have a mechanism that increases the
concentration of CO2 at the RuBisCO enzyme site. This takes
place when the C4 acid (malic or aspartic acid) from the
mesophyll cell is broken down in the bundle sheath cells to
release CO2 (CO2 pumping), this results in increasing the
intracellular concentration of CO2 . In turn, this ensures that
the RuBisCO functions as a carboxylase minimising the
oxygenase activity.
• In addition, in C4 plants, site of O2 evolution (mesophyll cell)
and site of RuBisCO activity (Bundle sheath cell) are different.
 C4 Pathway
• The evolution of the C4 photosynthetic system is probably one of
the strategies for maximising the availability of CO2 while
minimising water loss. C4 plants are twice as efficient as C3 plants
in terms of fixing carbon (making sugar). However, a C4 plant loses
only half as much water as a C3 plant for the same amount of CO2
fixed.
CAM (Crassulacean Acid Metabolism) Pathway/Dark CO2 Fixation
• CAM pathway was discovered by Oleary and Rouhani.
• They observed that CO2 fixation occurs during night in
members of Crassulaceae family (succulent xerophytes). Eg.
Kalanchoe, Bryophyllum, Opuntia, Agave, Aloe, Euphorbia,
Pineapple, Welwitschia (Gymnosperm)
• Succulents or CAM plants are characterised by
scotoactive stomata (stomata open during night and
remain closed during day time).
• Primary acceptor of CO2 = Phosphoenol pyruvate (PEP)
• First stable product = Oxaloacetic acid (OAA)
Mechanism of CAM Pathway
Photorespiration/Photosynthetic Carbon Oxydation (PCO) Cycle/ C2
Cycle/Glycolate Mechanism
• The light dependent uptake of O2 & release of CO2 in green
cells of C3 plants are called Photorespiration. This process
Photorespiration /Photosynthetic
createcarbon oxidation(PCO)
an important cycle/C
difference between Cycle/Glycolate.
C3 2and C4 plant.
It occursMetabolism
in chloroplast, peroxisome & mitochondria.
•
Conditions for photorespiration –
•
High light intensity (High O2, Low CO2) and
•
High temperature
•
• RuBisCO is characterized by the fact that its active site can
bind to both CO2 and O2 – hence the name. This binding is
competitive. It is the relative concentration of O2 and CO2 that
determines which of the two will bind to the enzyme. (Usually
RuBisCO has a much greater affinity for CO2 than for O2 ).
 Mechanism of Photorespiration
• In the photorespiratory
pathway there is neither
synthesis of sugars, nor of
ATP and NADPH2 rather it
results in the release of CO2
with the utilisation of ATP.
• Therefore, photorespiration is
a wasteful process.
• The biological function of
photorespiration is not known
yet.
 Warburg’s Effect
• The Warburg's effect is the decrease in the rate of photosynthesis by
high O2 concentrations.
• O2 is a competitive inhibitor of the CO2 fixation by RuBisCO. Furthermore,
oxygen promotes photorespiration which reduces photosynthetic output

Difference between bacterial and plant photosynthesis

Plant photosynthesis Bacterial photosynthesis
Pigment containing structures are Pigment containing structures are
1. 1.
thylakoids inside chloroplast chromatophores
Pigments are chlorophylls and Pigments are bacteriochlorophyll and
2. 2.
carotenoids bacterioviridin
It is oxygenic because PS II is present It is anoxygenic because PS II is absent (O2
3. which can photolyse the H2O. (O2 evolved) 3. evolution is absent)

Two pigment system PS I (P700) & PS II Only one pigment system is present whose
4. 4. photocentre B890. (PS II is absent)
(P680) are present.

5. Action spectrum is blue-red. 5. Action spectrum is infra-red

6. During light reaction NADP+ being 6. During light reaction NAD+ being reduced to
reduced to NADPH NADH
Factors Affecting Photosynthesis
• The rate of photosynthesis is very important in determining the yield of
plants including crop plants.
• Photosynthesis is under the influence of several factors, both internal
(plant) and external.
Internal (Plant) Factors :- External factors include :-
• The plant factors include the I. The availability of sunlight
I. Number of leaves II. Temperature
II. Size of leaves III. CO2 concentration
III. Age of leaves
IV. Water
IV. Orientation of leaves
V. Mesophyll cells and chloroplasts
VI. Internal CO2 concentration
VII. The amount of chlorophyll.
 The plant or internal factors are dependent on the genetic
predisposition and the growth of the plant.
Blackman’s Law of Limiting Factors
• When several factors affect any bio chemical process,
Blackman’s (1905) Law of Limiting Factors comes into effect.
This states the following:
• “If a chemical process is 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.”
• For example, despite the presence of a green leaf and optimal
light and CO2 conditions, the plant may not photosynthesize if
the temperature is very low. This leaf, if given the optimal
temperature, will start photosynthesizing.
External Factors (Light)
1. Light quality :
• Plant capture and utilised sunlight in the range of
400 – 700 nm (PAR).
• Most effective photosynthesis occurs in blue and red light.
• Least effective in green light.
2. Duration of light :
• Affects total production but does not affects the rate of
photosynthesis (if wavelength and intensity of light are
constant)
 External Factors (Light)
3. Light Intensity :-
• There is a linear relationship between incident light and CO2
fixation rates at low light intensities.
• At higher light intensities, gradually the rate does not show further
increase as other factors become limiting.
• Increase in incident light
beyond a point cause the
breakdown of chlorophyll and
decrease in photosynthesis.
• Light saturation occurs at 10%
of the full sunlight.
• Hence, except for plants in
shade or in dense forest, light
is rarely a limiting factor in
nature.
External Factors (CO2 Concentration)
• Carbon dioxide is the major limiting factor for photosynthesis.
• The concentration of CO2 is very low in the atmosphere
(between 0.03 and 0.04 percent).
• Increase in concentration up to 0.05 percent can cause an
increase in CO2 fixation rate, beyond this the levels can
become damaging over longer periods.
External Factors (CO2 Concentration)
• The C3 and C4 plants respond differently to CO2 concentrations.
At low light conditions, neither group responds to high CO2
conditions.
• At high light conditions, both C3 and C4 plants show increase in
the rates of photosynthesis.
• C4 -Plants show saturation at about 360 μl/L (0.036% or 360
ppm) while C3 responds to increased CO2 concentration and
saturation is seen only beyond 450 μl/L (0.045% or 450 ppm).
• Thus, current availability of CO2 levels is limiting to the C3
plants not for C4 plants.
External Factors (CO2 Concentration)
• The fact that C3 plants respond to higher CO2 concentration by
showing increased rates of photosynthesis leading to higher
productivity has been used for some greenhouse crops such as
tomatoes and bell pepper. They are allowed to grow in CO2
enriched atmosphere that leads to higher yields (CO2
fertilizing effect).
External Factors (CO2 Concentration)
• Atmospheric CO2 is not limiting factor for C4 plants and submerged
hydrophytes.
• CO2 compensation point :
• It is the point where rate of photosynthesis become equal to the rate
of respiration.
• At compensation point,
• Net productivity of the plant becomes zero and
• The plant do not show gaseous exchange with the atmosphere

• CO2 compensation point for C3 plants is 25-100 ppm.

• CO compensation point for C plants is 0-10 ppm.
External Factors (Temperature)
• The dark reactions being enzymatic are temperature
controlled. Though the light reactions are also temperature
sensitive they are affected to a much lesser extent.
• The C4 plants respond to higher temperatures (30° – 40° C)
and show higher rate of photosynthesis while C3 plants have a
much lower temperature optimum (20° – 25° C).
• The temperature optimum for photosynthesis of different
plants also depends on the habitat that they are adapted to.
Tropical plants have a higher temperature optimum than the
plants adapted to temperate climates.
External Factors (Water)
• Even though water is one of the reactants in the light
reaction, the effect of water as a factor is more through its
effect on the plant, rather than directly on photosynthesis.
• Water stress causes the stomata to close hence reducing
the CO2 availability.

• Besides, water stress also causes wilting of leaf, thus,

reducing the surface area of the leaves and their
metabolic activity as well.
External Factors (Inhibitor)
• Inhibitors are used as weedicides or herbicides.
• DCMU (Dichlorophenyl Dimethyl urea) / Diuron, CMU /
Monouron and PAN (peroxy acetyl nitrates) inhibit
photosynthesis by blocking PS-II as they stop electron
flow between P 680 and PQ.

• Diquat, Paraquat (violagen dyes) inhibit cyclic

photophosphorylation by blocking PS-I as they stop
electron flow between P 700 and Fd.