PHOTOSYNTHESIS
PHOTOSYNTHESIS
Photosynthesis is a Physico – chemical
process by which green plants use light
energy to synthesize Organic compounds.
Photosynthesis is important for two
reasons:
i) It is the primary source of food for all living
organisms.
ii) It is the only process responsible for the
release of Oxygen gas into the atmosphere
by plants.
EARLY EXPERIMENTS:
i) Joseph Priestley (1733- 1804):
He performed a series of experiments
(1770), that revealed the essential
role of air in the growth of plants.
He observed that a candle a burning
in a closed space (say a bell jar),
gets extinguished in a short time.
Similarly, a mouse kept in a closed
space, say a bell jar, would suffocate
soon.
He concluded that, a burning candle
or an animal that breathes the air, somehow damage the air.
But, when a mint plant is placed in the same bell jar,
it was observed that the mouse stayed alive and the candle
continued to burn.
So Priestley hypothesized that plants restore to the air,
whatever breathing animals and burning candles remove.
ii) Jan Ingenhousz (1730- 1799):
He conducted a similar experiment with an aquatic
plant, one kept in bright sunlight and the other in the
dark.
He showed that in the plant that is kept in the light,
small bubbles were formed around the green parts,
while no bubbles appeared in the plant kept in the
dark.
Later these bubbles were identified as Oxygen gas
bubbles.
Thus he proved that only green parts of the plants
could release Oxygen.
iii) Julius Von Sachs:
He showed that the green substance (chlorophyll) is
located in special bodies (chloroplasts).
He provided evidence (1854) for the production of
Glucose when plants grow and glucose is stored in
the form of starch.
He found that the green parts of the plants are the
parts where glucose is made.
iv) Englemann (1843- 1909):
He split the light using prism into its
component parts and illuminated a green
alga,
Cladophora placed in a suspension of
aerobic bacteria.
The bacteria were used to detect the sites of
Oxygen evolution.
He observed that, the bacteria accumulated
in the region of blue and red light of the
spectrum.
He first described the Action Spectrum of
photosynthesis, the Action Spectrum
resembles roughly the Absorption Spectrum.
v) Cornelius Van Neil (1897- 1985):
Based on his experiments with purple and green bacteria, he
made a contribution, a milestone, in the understanding of
photosynthesis.
He demonstrated that photosynthesis is essentially a light-
dependent reaction in which Hydrogen From a Hydrogen- donor
reduces Carbon dioxide to Carbohydrates.
The present day chemical equation for photosynthesis has been
given by him.
light
2H2A + CO2 ------------------- 2A + CH2O + H2O
In green plants water (H2O) is the Hydrogen donor and it is
oxidized to Oxygen.
Purple and green Sulphur bacteria use H2S as the
Hydrogen donor and so the oxidation product is Sulphur and
no Oxygen is produced.
This was later proved by using radioactive isotopes of Oxygen (
H218O) .
The overall correct equation for Photosynthesis is as follows.
Light
6CO2 + 12H2O --------------------- C6H12O6 + 6H2O + 6O2
SITE OF PHOTOSYNTHESIS:
Photosynthesis takes place mainly in the in the Mesophyll
tissue of the green leaves and to a small extent in the green
stems and floral parts ( sepals).
It takes place in the cell organelles called Chloroplasts.
Chloroplasts are present in large number in the Mesophyll
tissue in the leaves.
The chloroplasts are arranged in such a way that they get
maximum quantity of sunlight .
Under low intensity of light the chloroplasts are arranged
parallel to the walls and when the intensity is high they
become perpendicular to the walls of the cells.
Each chloroplast has two distinct regions. They are Stroma
and Grana .
Grana consists of stacks of Thylakoid membranes which
contain well arranged Chlorophyll and other pigment
molecules. Grana are the sites for the Light Reaction or
Photo- chemical Phase, where synthesis of ATP and
NADPH (ReducedNicotinamide Adenine Diphosphate)occurs.
Stroma has the enzymes for the Dark Reaction or
Biosynthetic Phase. Here reduction of CO2 into
Carbohydrates and formation of sugars occurs.
Site for Photosynthesis (Chloroplast)
PIGMENTS INVOLVED IN PHOTOSYNTHESIS:
A chromatographic separation of the leaf pigments
shows that there are four pigments.
Chlorophyll a, Chlorophyll b, Carotene and
Xanthophyll.
Chlorophyll a is bright green or bluish green.
Chlorophyll b is yellowish green .
Xanthophyll is yellow and Carotene is yellow, yellow-
orange or reddish.
Chlorophyll a is the main pigment associated with
photosynthesis, while Chlorophyll b, and
Carotenoids act as accessory pigments.
The accessory pigments help in two ways:
(i) They absorb light at different wavelengths and
pass it on to the Reaction Centre (Chlorophyll a).
(ii) They also protect the chlorophyll from Photo-
oxidation.
Differences Between Chlorophyll a & Chlorophyll b
Chlorophylls absorb light at the blue and red
regions.
The Absorption spectrum and Action spectrum
coincide closely showing that
photosynthesis is maximum at the blue and
red regions of the spectrum.
Photon:- Light is a form of energy and it travels
as a stream of tiny particles.
This is called as photon.
Quantum: The energy possessed by an individual
photon is called as quantum.
The energy of photon depends upon frequency of
the wavelength of radiation.
Blue light provides more energy than red light by
utilizing one photon because
blue light has higher frequency than red light.
(i) Absorption spectrum:
o All the photosynthetic pigments do not absorb
the same wavelengths of the visible spectrum of
light.
A curve obtained by plotting the amount of
light absorbed by a particular pigment against
the different wavelength of light is
Absorption Spectrum.
(ii) Action spectrum:
It is the curve showing the actual rate of O2
evolved or CO2 consumed during photosynthesis
against the different wave lengths of light
absorbed by the same pigment.
Absorption and Action spectrums For Chl a & Chl b
PHOTO-SYSTEMS:
The pigments are organized in the Thylakoid
membranes into two discrete Photo-systems, Photo-
system I (PS I) and Photo-system II (PS II).
Each photo-system has one specific Chlorophyll a and
many other pigments bound by proteins.
The Chlorophyll a forms the Reaction Centre,
where the actual reactions take place, while the other
pigment molecules form the
Light Harvesting Complex (LHC), also called as
Antennae.
In PS I, the reaction centre is P700, which is a
Chlorophyll a molecule that absorbs light at 700 nm.
In PS II, the reaction centre is P680, which is a
Chlorophyll a molecule that absorbs light at 680 nm.
Reaction Centre( Light Harvesting Zone)
Difference Between Photo Systems I & II
MECHANISM OF PHOTOSYNTHESIS:
Photosynthesis occurs in two steps:
(i) Light reaction or Photochemical phase.
(ii) Dark reaction or Biosynthetic phase.
Photochemical phase directly depends on light, the
pigments absorb the light energy and convert it into ATP.
During this phase, Water is split (photolysis), Oxygen is
liberated and NADPH is formed.
ATP and NADPH formed in this phase are used up during
the Biosynthetic phase. Electron transport (ATP
synthesis) and Photolysis the two major events in light
reaction.
Biosynthetic phase does not directly depend on light, but
depends on the products of light reaction i.e. ATP and
NADPH.
The biosynthetic process continues for sometime after the
light becomes unavailable but soon stops.
In this phase, Carbon dioxide is reduced to
Carbohydrate, i.e, Glucose is synthesized.
PHOTOLYSIS OF WATER:
In This process water is split intp Protons,
Electrons and Oxygen.
2H2O--------------------------- 4H+ + 4 e- + O2
The water splitting complex is associated with
Photosystem II, which is located on the innerside
of the Thylakoid membrane.
The electrons are used to replace the electrons lost
by PS II.
Manganese, Chloride and Calcium ions play
important roles in photolysis of water.
The Oxygen liberated is liberated into the
atmosphere.
The protons are used for reducing NADP to
NADPH.
PHOTOPHOSPHORYLATION:
It is the process of formation of ATP in the
chloroplasts, making use of the light energy.
The protons formed during photolysis of water
accumulate inside the Thylakoid membrane
resulting in a proton gradient.
The energy released by the protons, when they
diffuse across the Thylakoid membrane into the
Stroma following the H+ concentration gradient, is
used for the formation of ATP.
It occurs in two ways :
(i) Non-cyclic Photo- Phosphorylation and
(ii) Cyclic Photo- Phosphorylation.
i) Cyclic Photophosphorylation:
When PS I (P700) is activated by light, it is
raised to high energy level and emits a pair of
electrons.
These electrons are captured by the Primary
acceptor, which passes them on to Ferridoxin,
Plastoquinone, Cytochrome complex and
Plastocyanin.
Finally the de-energized electrons come back to
P700.
Since the electrons come back to P700 , this
process of ATP formation is known as
Cyclic Photophosphorylation.
Cyclic Photophosphorylation
ii) Non-cyclic Photophosphorylation:
When P680 (PS II) acquires sufficient amount of light
energy, it becomes excited and emits electrons.
The electrons (with high energy) run down an
electron transport chain, consisting of a Primary
acceptor, Plastoquinone, Cytochrome complex
and Plastocyanin to P700 which transfers
them to Ferredoxin.
Ferredoxin in turn transfers the electrons to
NADP, which along with the protons is reduced to
NADPH.
The electrons lost by PS II are replaced by the
electrons generated by splitting of water.
Since the electrons lost by PS II do not come
back to it, this process of ATP formation is
called Non-cyclic Photophosphorylation.
Non Cyclic Photophosphorylation
Differences Between Cyclic and Non-
cyclic Photophosphorylation
CHEMI-OSMOTIC HYPOTHESIS:
Chemi-osmotic hypothesis has been put forward to
explain the mechanism of ATP synthesis.
ATP synthesis is linked to the development of a
proton gradient across the membranes of
Thylakoids.
When electrons are transported through the
electron transport system (ETS), the Protons
accumulate inside the Thylakoid membranes.
Since the splitting of the water molecules or
photolysis takes place on the inner side of the
Thylakoid membrane, the Protons produced,
accumulate within the lumen of the Thylakoids.
As the electrons are passed through Photosystems,
protons are transported across the membrane.
This happens because protons are removed from the Stroma
for the following reasons:
(i) The primary electron acceptor is located towards the outer
side of the membrane and transfers its electrons to the H
carrier, so this molecule removes a portion from the Stroma
while transporting an electron and releases it into the lumen
or inner side of the membrane.
(ii) The enzyme NADP – reductase is located on the Stroma
side of the membrane. Along with the electrons coming from
the PS I, protons are also needed to reduce NADP and so
these protons are also removed from the Stroma.
As a result, the Protons in the Stroma decrease in number,
while the protons in the lumen, increase in number, thus
creating a proton gradient across the Thylakoid membrane.
This gradient is important because energy is released by the
breakdown of this gradient.
The gradient is broken down due t the movement of protons
across the membrane through trans-membrane channel of the
Fo of the ATP synthetase; the other portion of ATP synthase
called F1 undergoes conformational changes with the energy
provided by the breakdown of proton gradient and synthesizes
several molecules of ATP.
Thus Chemi-osmosis requires:
a membrane,
a proton pump,
a proton gradient and
ATP synthetase.
Energy is used to pump Protons across the
membrane, to create a gradient or high
concentration of protons within the lumen of the
Thylakoid.
ATP synthetase has a channel that allows the
diffusion of protons back across the membrane
which Releases energy.
This energy activates ATP synthetase enzyme that
catalyses the formation of ATP.
ATP Synthesis Through Chemiosmosis
ATP Synthesis Through Chemi-osmosis
BIOSYNTHETIC PHASE (DARK REACTION):
The process by which carbon dioxide is reduced to
carbohydrates is called carbon fixation.
The process makes use of the ATP and NADPH
produced in the photochemical (light reaction) phase.
Carbon fixation occurs in the Stroma of chloroplasts, by
a series of enzyme- catalyzed reactions.
C3 PATHWAY OR CALVIN CYCLE:
The path of carbon in the dark reaction was traced by
Melvin Calvin through a technique called
Autoradiography, using 14C, hence this pathway is
termed as Calvin cycle.
Calvin cycle consists of three phases viz:
(i) Carboxylation
(ii) Reduction
(iii) Regeneration of RuBP.
Calvin Cycle (Dark Reaction)
Calvin cycle consists of three phases viz:
(i) Carboxylation
(ii) Reduction
(iii) Regeneration of RuBP.
(i) Carboxylation:
Six molecules or Ribulose 1,5 bisphosphate react
with six molecules of carbon dioxide to form six
molecules of a short lived 6C-compound.
The reaction is catalysed by RuBP-carboxylase
(Rubisco)
The six molecules of the 6C-compound break into 12
molecules of 3-phosphoglyceric Acid (PGA), a 3C –
compound.
PGA is the first stable compound in this pathway.
(ii) Reduction:
12 molecules of Phosphoglyceric acid are converted
into 12 molecules of 1,3- di-phosphoglycerate and
then reduced to Phosphoglyceraldehyde (PGAL)
using ATP and NADPH molecules respectively
Two molecules of PGAL are diverted for the
synthesis of sugar and then starch.
(iii) Regeneration RuBP:
For the cycle to continue, the primary acceptor of
carbon dioxide i.e. RuBP has to be regenerated.
10 molecules of PGAL, by a series of complex
reactions are converted into 6 molecules of 5C-
compound, RuBp.
The formation of y molecules of RuBP requires six
ATP molecules.
PHOTORESPIRATION:
In C3 plants, RuBP-carboxlyase, which is the
main enzyme of photosynthesis functions as
Oxygenase at high temperatures and high
Oxygen concentration.
It catalyses the oxidation of RuBP into one
molecule of a 3C – compound, Phosphoglycerate
and one molecule of a 2C-compound,
Phosphoglycolate.
Phosphoglycolate is quickly converted into
Glycolate and transported to Peroxysomes.
In the Peroxisome, Glycolate is oxidized to
Glyoxylate and is converted into amino acid
Glycine
Glycine enters the Mitochondria and two
Glycine molecules give rise to one molecule of
Serine and one molecule of Carbon dioxide.
Serine is picked up by Peroxysomes and is
converted to Glycerate by a series of reactions.
The Glycerate leaves the Peroxisome and enters
the chloroplast, where it is phosphorylated to form
PGA.
The PGA enters Calvin cycle and thus 75% of the
cabon lost during oxygenation of RuBP is recovered,
but 25% is lost, as one molecule of carbon dioxide is
released.
Photorespiration is a wasteful process:
There is no sugar or ATP formation.
CO2 is released and ATP consumed instead of
being formed as in case of Respiration.
Photo- respiration
Anatomy of C4 plants:
The C4 plants have two types of photosynthetic cells
– Mesophyll cells and Bundle sheath cells. The
bundle sheath cells are arranged in a wreathlike
manner.
This kind of arrangement is called Kranz anatomy.
The C4 plants bear dimorphic chloroplasts – the
chloroplasts in Mesophyll cells are Granal and
in Bundle sheath cells are Agranal. The Granal
chloroplasts contain Thylakoids that are stacked to
form Grana (as in C3 plants), but in Agranal
chloroplast, the Grana are absent and the
Thylakoids are present only as Stroma lamellae.
These two types of cells allow the occurrence of light
reactions and carbon reactions separately in each
type.
In C4 plants, light reactions occur in
Mesophyll cells and CO2 assimilation is
carried out in Bundle sheath cells. This
type of cellular arrangement does not allow
O2 released in Mesophy11 cells to escape
to Bundle sheath cells.
o Thus, Rubisco that oxidation of RuBP is
completely prevented. Moreover, C4 plants
have CO2 concentrating mechanism where
the concentration of CO2 remains high in
the vicinity of Rubisco in the bundle sheath
cells. This favours Carboxylation and
suppresses photorespiration.
Monocots Leaves With Kranz Anatomy
C4 PATHWAY OR HATCH-SLACK PATHWAY:
This pathway of carbon fixation occurs in plants
like Maize, Sugar cane, Pearl millet and
Amaranthus.
The plants which employ this mechanism of
Carbon fixation show the presence of two types of
photosynthetic cells i.e. Mesophyll cells and
Bundle sheath cells (Kranz anatomy)
The chloroplasts are dimorphic i.e., those in
Mesophyll cells are Granal and those in the Bundle
Sheath cells are Agranal.
Phospho-Enol-Pyruvate (PEP) is the Carbon
dioxide acceptor, present in the Mesophyll cells and
the reaction is catalyzed by the enzyme Phospho-
Enol-Pyruvate Carboxylase (PEP-case).
The first stable product is Oxalo Acetic Acid
(OAA), which is a 4C-compound.
OAA is converted into Malic acid and transported
to Bundle sheath cells, where it is decarboxylated
into Pyruvic acid.
The Carbon dioxide liberated is used for Calvin
cycle in the bundle sheath cells.
Pyruvic acid is transported back to Mesophyll cell,
where it is used for regeneration of PEP, utilizing
two molecules of ATP for each pyruvate.
It is an adaptive mechanism to avoid
photorespiration, the Decarboxylation of Malic acid
in the Bundle sheath cells maintains a high
concentration of CO2 Rubisco, so that it functions
as Carboxylase only and not as Oxygenase.
Mechanism of C4 Pathway
Differences Between C3 and C4 Pathways
FACTORAS AFFECTING PHOTOSYNTHESIS:
Blackman’s Law of Limiting Factors
When a physiological process is controlled by a number of
factors, the rate of the reaction depends on the slowest
factor. This means that at a given time, only the factor
which is the least (limiting) among all the factors, will
determine the rate of the reaction.
External factors:
(i) Light:
Light quality and light intensity influence photosynthesis.
Light of wavelength between 400 nm and 700nm is
effective for photosynthesis and this light is known as
photosynthetically active radiation. (PAR)
As the intensity of light increases the rate of
photosynthesis increases. But at higher light intensities,
the rate of photosynthesis decreases; it may be due to two
reasons :
1. Other factors needed for photosynthesis may be limiting.
2. Destruction (Photo-oxidation) of chlorophyll.
(ii) Temperature:
The photochemical phase remains unaffected by
temperature. But the biosynthetic phase that
involves enzyme- catalyzed reactions is sensitive to
Temperature, it may be due to the following
reasons:
1. At high temperatures, the enzymes become
denatured (inactive)
2. At low temperatures, the enzymes may be
inactivated.
(iii) Carbon dioxide concentration:
In C3 plants, the rate of photosynthesis increases
with increase in CO2 concentration, upto about
500ul.l-1.
In C4 plants, the saturation is reached at a
concentration of about 360 ul.l-1
Blackman’s Law of Limiting Factors
(iv) Available soil water:
Water influences photosynthesis in two ways:
1. If available water decreases and plants show water
stress, the stomata will not open and hence there will
be a decreased supply of Carbon dioxide.
2. Reduced water potential in the leaf reduces leaf
expansion and reduction in the photosynthetic surface
area.
Role of RUBISCO in C3 and C4 Photosynthesis:
Role – RuBP carboxylase in main enzyme, which brings
about Carboxylation during Calvin cycle in
photosynthesis.
It combines CO2 with RuBP and forms the 3-
phosphoglyceric acid. Under high concentration of
Oxygen and with increase of temperature, the enzyme
RuBP Carboxylase acts as RuBP Oxygenase and its
affinity for CO2 decreases and for O2 increases. Thus,
more and more photosynthetically fixed carbon is lost
(photorespiration).
BIBLIOGRAPHY
Ch: 13 ‘Photosynthesis’ from NCERT Text Book of
Biology Class XI.
Ch: 13 ‘Photosynthesis’ Modern’s ‘abc’ Biology.
Class XI.
Ch: 13 ‘Photosynthesis’ Pradeeps’s A Text Book of
Biology, Class XI.
‘Ch: 13 ‘Photosynthesis’ Together with’ Biology,
Class XI, Rachna Sagar.
Ch: 13 ‘Photosynthesis’ from the printed noted
prepared by Ms A. Jayashree DPS Hyderabad.
Pictures from Google Chrome.
