Plant Water Relations

Water Molecule
Water is a dihydride of oxygen. Both the hydrogen atoms are attached to oxygen by covalent
bonds.
Due to sharing of electrons, there is a displacement of hydrogen atoms towards oxygen
atom. As a result, the angle between the two hydrogen atoms comes to be I04-5°C
The oxygen atom develops a partial negative charge (5 —) while each of the two
hydrogen atoms comes to have a partial positive charge (8+).
The molecules which show charge separation, as found in water between hydrogen and
oxygen atoms are called polar molecules.
➢ Ice occupies more space due to the space occupied by hydrogen
bonds. Water is heaviest at 4°C. Because of this, ice floats over the
surface of water.
➢ Inspite of its being weak, hydrogen bonds increase the energy
required to separate water molecules from solid state to liquid state or
from liquid state to vapour state. This raises melting point, boiling point
and latent heat of vaporisation of water. Water possesses high surface
tension and cohesiveness due to the loose hydrogen bonding between
water molecules.
➢ These peculiar properties have made water liquid between nearly 0°
—100°C. The liquid nature of water allowed dissolution, reactivity and
development of chemicals that paved the way for origin of life. Water is
also transparent. It allows light to pass through it. This is essential for
aquatic life. Water also possesses the maximum solvent power for any
liquid. It allows dissolution of electrolytes and polar non-electrolytes.
Further, it forms shells around ions and other solutes.
 Importance of Water to Plants
➢cells.Water is the most abundant constituent of living matter, 70—80% of
➢protoplasm.
It is dispersion medium of crystalline colloidal complex of

➢configuration
Proteins and nucleic acids develop their three dimensional
and reactivity in the presence of water.
➢the basic
Phospholipids form abilayer over its surface. Phospholipid bilayer is
framework of all cell membranes.
➢ It is the general solvent of living beings.
➢Substances enter the cells mostly dissolved in water.
➢Osmolarity is expressed only in water.
➢It helps cells and their organelles to maintain turgidity.
➢Water is essential for cell growth.
➢changes
Many movements of the living world occur due to turgor
caused by gain or loss of water, e.g., opening and closing of
guard cells, folding and unfolding of leaves, opening and closing of
flowers, movements of Sensitive Plant.
➢ It is a medium of transport.
➢ It has a high force of cohesion and adhesion. This is
helpful in holding water molecules in the liquid state, ascent of
sap and wetting of body parts.
➢ Water is required for seed germination.
➢ It has a high thermal conductivity and high latent heat.
The two properties spread heat quickly and function as buffer
against sudden temperature changes in living beings.
➢ It causes dissociation of electrolytes into ions.
➢ Most biochemical reactions occur with water as solvent.
➢powerWater is a reactant in photosynthesis. It produces reducing
and evolves oxygen.
➢ Water is an end product of respiration.
➢molecules
It takes part in breakdown of larger molecules into smaller
through a process called hydrolytic splitting.
➢ (C6H10O5)„ + n H20-> n C6H1206
 Absorption and Movement of Water
Diffusion
The movement of the molecules from the region
of higher concentration to the region of lower
concentration is known as diffusion.
Osmosis
The movement of solvent molecules from the region
of their higher concentration to the region of their
lower concentration through a semi permeable
membrane is called osmosis.
Diffusion of Copper Sulphate in Water
➢
Take a beaker.
➢ Fill a beaker with water.
➢
Add a few crystals of copper sulphate.
➢ Water surrounding the crystals turns dark blue.
➢
The water in the beaker turns
light blue in colour (copper suplhate
crystals completely dissolve in water).
➢ Diffusion of copper sulphate molecules is due to the reason that

molecules are in a state of constant motion (kinetic energy).

 Osmosis
The movement of solvent molecules from the region
of their higher concentration to the region of their
lower concentration through a semi permeable
membrane is called osmosis.
Demonstration of Osmosis Using Potato Osmoscope
➢ Peel off the skin of a large sized potato with a scalpel.
➢ Cut its one end to make the base flat.
➢ Make a hollow cavity in the potato almost up to the bottom.

➢ Add sugar solution into the cavity and mark the level by
inserting a pin in the wall of the cavity of the tuber.
➢ Place the potato in the beaker containing water.

➢ After sometime, it will be noticed that the level of solution in the

cavity rises. This is due to the phenomenon of osmosis.
➢ This experiment demonstrates that living cells of potato act as
differentially permeable membrane.

➢ Water molecules from a region of their higher concentration

(water in the beaker) move into a region of their lower concentration
(sugar solution in potato cavity) through the differentially permeable
cell membrane of the potato cells.
 Importance of Osmosis in Plants
➢ In the absorption of water by plants

➢ Cell to cell movement of water occurs throughout the plant

body

➢
The rigidity of plant organs is maintained
➢ Leaves become turgid and expand due to osmotic pressure
➢
Growing points of root remain turgid
and penetrate the soil particles
➢
The resistance of plants to drought and
frost ➢Movement of plants and plant
parts ➢Opening and closing of stomata
Diagram showing relationship of Osmotic Pressure (OP),
Turgor Pressure (TP) and Wall Pressure (WP)
 Plasmolysis
➢ If a plant cell is placed in a highly concentrated sugar or salt
solution (i.e., hypertonic solution), water from the cell sap flows
out due to exosmosis through the plasma membrane outside the
cell.

➢ The loss of water from the cell sap causes contraction

or shrinkage of the protoplast. Since the cell wall is firm
and less elastic, it cannot keep pace with the contraction of
plasma membrane.
➢ Ultimately the protoplasm separates from the cell wall and
contracts or shrinks to a spherical shape. This condition is called
plasmolysis.
A. Normal Cell B. Incipient Plasmolysis C. A
Plasmolysed Cell
Various Stages in Plasmolysis
 Incipient Plasmolysis
The stage of plasmolysis at which
➢

the first sign of shrinkage of cell contents from

cell wall becomes detectable is called incipient plasmolysis.
Evident Plasmolysis
The stage when the cell wall
➢
has reached its limit of contraction
and cytoplasm has detached from cell wall attaining
spherical shape is called evident plasmolysis
 Absorption of Water by Plants
➢ Plants absorb water through the entire surface - roots, stems and
leaves. However, mainly the water is absorbed by roots. The area of
young roots where most absorption takes place is the root hair zone.
The root hairs are delicate structures which get continuously replaced
by new ones at an average rate of 100 millions per day. The root hairs
lack cuticle and provide a large surface area. They are extensions of the
epidermal cells. They have sticky walls by which they adhere tightly to
soil particles. As the root hairs are extremely thin and large in number,
they provide enormous surface area for absorption. They take in water
from the intervening spaces mainly by osmosis.
➢ Water in the roots move by two pathways. They can be classified
as1) Apoplast pathway2) Symplast pathway
Areas of Root Involved in Absorption and Translocation of Water
 Mechanism of Water Absorption
➢ Water can be absorbed by two methods:
➢ Active absorption
➢ Passive absorption
Active Absorption
➢ Water is absorbed due to activities going on in roots. Absorption of
water occurs with the help of energy in the form of ATP, which is released
due to metabolic activities of root cells such as respiration. Absorption
takes place against concentration gradient - even when the concentration
of cell sap is lower than that of soil water.
Passive Absorption
➢ Passive absorption is by osmosis. Passive absorption takes place along the
concentration gradient - when the concentration of cell sap is higher than that
of soil water. Water is absorbed when transpiration rate is high or soil is dry.
Due to high transpiration rate, water deficit is created in transpiring cells.
➢ Rapid transpiration removes water and reduces turgor pressure in
living cells of root. The suction force thus developed is transmitted to root
xylem. It pulls water from surrounding root cells to make up water deficit.
 Ascent of Sap
The upward movement of water through stem is called
ascent of sap
Path of ascent of sap
➢
The water absorbed by the root
hairs is moves upwards via root tissues (cortex,
endodermis, and pericycle) and finally enters the xylem.
➢ After entrance it starts its upward movement until it reaches
mesophyll of leaves. Bulk of water enters mesophyll cells and
finally evaporates through stomata.
➢
Only a small amount of water is utilized
by plants for metabolism, rest is lost in transpiration.
• Fig1. Ascent of sap

Fig:- Ascent of sap

 Mechanism of Ascent of Sap
A number of theories have been explain about
the mechanism of ascent of sap
➢ Root pressure theory
➢ Vital theories
➢ Transpiration pull theory
Root pressure theory
➢ In, 1956 Stocking defined root pressure as “Pressure developing in
tracheary elements of xylem as a result of metabolic activities of
roots”.
➢ If a well watered plant is cut near its base, the xylem sap is seen
to flow out through cut end with a pressure. This pressure is actually
the hydrostatic pressure developed in the root system called root
pressure.
 Vital theories
➢ According to vital theories living cells are required for ascent
of sap. These have been proposed by many workers namely
Godlewski (relay pump theory), Bose (pulsation theory) and
Molish.
➢ These theories were discarded because it was discovered
that some poisons like picric acid and carbolic acid can also be
translocated, thus fundamental basic of vital theories fails

Transpiration pull theory (Cohesion – tension

theory)
➢ In 1894, Dixon and Jolly proposed this theory which is based on
following features.
➢ Cohesion and adhesion properties of water molecule to
form unbroken continuous water column in xylem. b.
Transpiration pull or water tension is exerted on xylem
column
➢ According to this theory, evaporation of water from the cells of
leaf is responsible for raising water from the root. Evaporation results
in a reduced water potential in the cells next to the xylem. Water
therefore enters these cells from the xylem sap which has a higher
water potential, passing through the moist cellulose cell walls of the
xylem vessels at the ends of the vein.
➢ The xylem vessels are full of water and, as water leaves them, a
tension is set up in the columns of water. This is transmitted down the
stem all the way to root by cohesion of water (water molecules have
high cohesion) and they also tend to stick to the vessel walls, a force
called adhesion.
Transpiration-cohesion Theory
 Transpiration
➢ Transpiration is the loss of water from the aerial parts of the plant in
the form of water vapour. Transpiration may be of the following types:
➢ Stomatal
➢ Transpiration taking place through the stomata is called
stomatal transpiration.
➢ Cuticular
➢ The surface of the plant is covered by a thin layer of cuticle through
which water vapour is lost during transpiration. This is called cuticular
transpiration.
➢ Lenticular
➢ The process of loss of water vapour through the lenticels is called
lenticular transpiration.
➢ Of the three types, maximum transpiration is accounted
through the stomata.
 Significance of Transpiration
Absorption of water
➢ Transpiration influences the rate of absorption of water from the soil.
Water movement
➢ By transpiration, water moves upwards and as it passes into the cell
vacuoles, it makes the cells turgid. This gives form and shape to cells and
plant as a whole.
Mineral salt transport
➢ The water stream moving upwards carries dissolved minerals with it.
Transpiration also helps in distributing these minerals throughout the plant.
Cooling
➢ The evaporation of water during transpiration cools the leaves.
Protection from heat injury
➢ Some plants like cacti, retain water by reducing transpiration. This
saves the plants from high temperatures and strong sunlight.
 Factors Affecting Rate of Transpiration
➢ Extern Temperature
➢ Higher the temperature more is the transpiration.
➢ Light
➢ Light causes stomata to open and hence increase the water loss from plant.
➢ Availability of soil water
➢absorption
When the soil gets dry, soil solution becomes more concentrated and the rate of
by cells decreases. This leads to reduction in transpiration and stomata
close quickly to keep the water loss to minimum.
➢ Atmospheric humidity
➢ High humidity means high water vapour pressure outside and it results in
lower rate of transpiration and as the humidity decreases rate of transpiration
increases.
➢ Wind
➢transpiration.
The wind removes water vapour and thus increases the rate of
High winds lead to stomatal closure to stop the rapid water loss
and hence bring a drop in rate of transpiration. Moderate winds may also
reduce transpiration by lowering the temperature of leaf.
➢ Atmospheric pressure
➢ Lower the atmospheric pressure, higher is the rate of transpiration.
 Stomata
➢ Stomata are openings, generally on the under surface of the leaves.
These openings are guarded by bean-shaped cells (dumb-bell shaped in
grasses) called the guard cells.
➢ The walls of the guard cells facing the stomata are thickened. Due to an
increase in the concentration of K+ ions inside the guard cells, the water
potential reduces. This results in water entering the guard cells and they
become turgid. The outer thin walls bulge which pulls the thicker inner walls
apart, opening the stomata.
➢ For closure the reverse occurs - loss of K + ions, increased water
potential, exit of water and the cells become flaccid. This brings the inner
walls close together and closes the stomata.
➢ Transpiration is maximum during the day, from morning to midday. This
is the time stomata remain open. They close during the evenings and at night
when the transpiration rate is low
➢ Transpiration is the loss of water in the form of water vapour from the
aerial parts of the plant.
 +
Diagram Showing Normal Responses of Stomata to Light, CO2, pH, K
ion and Water Deficiency
Water Movement through the leaf to the atmosphere in the form of vapour
 Role of Stomata in Transpiration
➢ Since most of the water (90%) is lost through stomata, plants
regulate the degree of stomatal opening to reduce the water loss.
Structure of Stomata
➢ Each stomata consists of a minute pore called stoma
surrounded by two guard cells.
➢ The stoma acts as a turgor operated valve which closes and opens
according to the turgidity of guard cells.
➢ The guard cells are the only epidermal cells that contain
chloroplasts. The guard cells have unevenly thickened walls.
➢ The cell wall around stoma is tough and flexible and the one away
from stoma is thinner.
➢ The shape of guard cells differs in dicots and monocots
though the mechanism remains the same.
 Mechanism of Stomatal Action
• The mechanism namely the opening and closing of stomata depends
upon the turgor pressure in the guard cells. When the guard cells
are turgid, the stoma opens and when the guard cells lose water,
stoma closes.

Stomatal Movement in Dicot Plants

Stomatal Movement in Monocot Plants
• Guttation
• The loss of extra water in liquid drops from margins of leaves of
herbaceous plants when root pressure is high and transpiration
is low.
• This process is very common during warm humid nights.
• This process occurs in plants growing in high soil moisture.
• It occurs through specialised pores called hydathodes present
near the vein endings.

Nasturtium Leaf Showing Guttation at the Margin of Leaf

 Imbibition

• Imbibition is the phenomenon of adsorption of water

by the solid particles of a substance without forming
a solution.
• Importance of Imbibition to the Plants
• Imbibition is the initial step in seed germination.
• Imbibition causes swelling of seeds and results in
the breaking of testa.
• Imbibition is dominant in the initial stage of
water absorption by roots.
• The water moves into ovules which are ripening
into seeds by imbibition.
 Photosynthesis
• The first photosynthetic organism probably appeared almost three billion
years ago. With the evolution of photosynthesis, however, organisms
began to change the face of our planet and, as a consequence, to exert
strong influences on each other. Organisms have continued to change the
environment, at an ever increasing rate, up to the present day.
• Primitive earth had reducing atmosphere (an-oxygenic). The atmosphere
in which the first cells evolved consisted largely of ammonia, water,
methane and other carbon gases. Gradually methane and ammonia were
used up in the formation of the organic molecules on which the first cells
were dependent. Their energy probably came from anaerobic glycolysis.
• Then, there slowly evolved photosynthetic organisms that used carbon
dioxide as their carbon source and released oxygen. As these photosynthetic
organisms multiplied, they provided a new supply of organic molecules and
free oxygen began to accumulate. Oxygen - consuming organisms have an
advantage over those that do not use oxygen and are active and complex.
• Life on earth continues to be dependent on photosynthesis both for its
oxygen and for its carbon containing fuel (food) molecules. Photosynthetic
organisms capture light energy and use it to form carbohydrates and free
oxygen from carbon dioxide and water, in a complex series of reactions.
• Photosynthesis is a process by which plants and other
organisms (like algae and some bacteria) synthesize their
own food in the presence of light.
Or
• Photosynthesis is a multi-stage process consisting of a series
of light requiring reactions and a series of chemical reactions.
• The former, collectively called the light reaction involves light
energy, absorbed by chlorophyll and carotenoids. This light
energy 'splits' water molecules (photolysis) into hydrogen and
oxygen. The latter is released as gaseous oxygen. In the non-
light dependent reactions, or dark reactions, hydrogen
combines with (reduces) carbon dioxide to form carbohydrate.
The overall equation can be represented as shown below:
 Photosynthetic Apparatus
• The photosynthetic pigments chlorophylls are present in the leaves
because of which the leaves are green in colour and are called the
photosynthetic organs. The leaves have the following structure:

Histological Details of a Leaf in Cross Section

• As shown in the diagram, the mesophyll region of the
leaves has cells made of Palisada tissue which contain
the pigments. Inside the cell the pigments are
present in organelles called chloroplasts which are
also known as the photosynthetic organelles.
• Chloroplasts can be seen easily under a light
microscope (4 6 micrometer in diameter). A
continuous double membrane surrounds the
chloroplast. In the chloroplast of higher plants, stacks
of lamellar structures, called grana (singular granum)
are present. Its inner membrane lines the lumen of the
chloroplast and it is called matrix or stroma. In the
cross-section of the grana, lamellae form sac like
structures called thylakoids. A thylakoid of one
granum is connected to other granum with the help of
a membrane. It is called stroma lamellae.
• Section of Chloroplast as seen under electron microscope
• The lower epidermal layer of the leaves have
specialised cells which guard the openings called the
stomata. It is through the stomata that the carbon
dioxide enters and the oxygen produced leaves the
plant body. Water is made available by the vascular
bundles of the midrib and the veins in the leaves.
From the vascular bundles, the water moves by
osmosis into the photosynthetic cells.
• In higher plants, the leaves along with the chloroplasts
are considered the photosynthetic apparatus of the
plant. Certain young regions may, however, contain
chlorophyll. However, in the lower plants like algae,
the entire plant body contains the pigments. The
exchange of gases and uptake of water takes place
through the general surface of the plant body.
 Photosynthetic Pigments
• Molecules of chlorophyll-a, chlorophyll-b, carotene and
xanthopyhll are situated in the thylakoid membranes.
• For light energy to be used by living systems, it must
first be absorbed. A pigment is any substance that
absorbs light. Chlorophyll, the pigment that makes
leaves green, absorbs light in the violet and blue
wavelengths and also in the red because it reflects
green light, it appears green. Different pigments
absorb light energy at different wavelengths. The
absorption pattern of a pigment is known as the
absorption spectrum. The absorption spectrum of
chlorophyll is between 400 nm and 700 nm. This
portion of the spectrum is known as photosynthetically
active region (PAR).
• Electromagnetic Spectrum: Action and Absorption Spectrum
of Chlorophyll a and b
The action spectrum shows how effective these pigments are in
stimulating photosynthesis.In plants, chlorophyll a is the pigment
directly involved in the transformation of light energy to
chemical energy.
 Structure of Chlorophyll
• Structurally all types of chlorophyll resemble one another. All of
them contain four pyrrole rings [also called porphyrin] which are
linked together by methane bridges (-CH=). The skeleton of each
pyrrole ring is made up of five atoms - four carbon and one nitrogen
with magnesium in the centre as nucleus. One pyrrole ring is
esterified with a long chain alcohol - phytol. This side chain-phytol
is long and is composed of insoluble carbon and hydrogen atoms
which helps to anchor the chlorophyll molecules with the
thylakoids.
• In plants, there are 2 types of chlorophyll - namely chlorophyll a and
b. Chlorophyll molecule looks like a tadpole with porphyrin head and
phytol tail. Chlorophyll a has methyl group (-CH3) at position and
aldehyde (CHO) group in chlorophyll b.
• Chlorophyll a is the major pigment involved in trapping light
energy and converting it in to electrical and chemical energy. It
acts as a reaction centre.
 th
• Chlorophyll b constitutes about 1/4 of the total
chlorophyll content. It acts as an accessory
pigment and helps broaden the spectrum of light
absorbed during photosynthesis. Chlorophyll b
absorbs a different wavelength of light other than
that absorbed by chlorophyll a. On absorbing light,
it becomes excited and transfer its to chlorophyll a
molecule.
• Another group of pigments are called carotenoids.
The carotenoids are red, orange or yellow
pigments. In the green leaf, their colour is masked
by the chlorophylls, which are more abundant.
• Carotenoids like chlorophyll are embedded in the
thylakoid membrane of the chloroplasts. They are
accessory pigments and harvest light from different
regions of the spectrum. The light captured by these
pigments are channelled it to the reaction centre,
where light energy is converted into electrical energy.
The chemical structure of chlorophyll X*:-CH3 in chlorophyll a, - CHO
in chlorophyll b
Conversion of Light into Electrical Energy
Colour and Form of Pigments That
Constitute Chlorophyll
Name of pigment Colour

Chlorophyll a Yellow - Green

Chlorophyll b Blue - Green

Carotene Orange

Xanthophyll Yellow
 Mechanism of Photosynthesis
Mechanism of Photosynthesis revolves around Light
and Dark reaction which as the name suggests are
light dependent and independent respectively.
The entire mechanism can be basically divided
into following stages:
➢ Light Reaction (Hill Reaction)
➢ Dark reaction (Calvin Benson Cycle)
 Light Reaction
First stage of Photosynthesis which converts light energy
into chemical energy in the form of ATP and NADPH.
Location: Thylakoid membrane inside choloroplast.
The thylakoid membrane contains some integral
membrane protein complexes which catalyze the light
reactions.
The following four complexes work together to create the
products ATP and NADPH:
➢ Photosystem I (PSI)
➢ Photosystem II (PSI)
➢ Cytochrome b6f complex
➢ ATP synthase
 Steps in Light Reaction
Steps in Light reaction occurs in following steps, together
popularly known as Z scheme of Light reaction:
➢ Cyclic Phosphorylation
Excited electrons from Photosystem I (P700) moves through a
series of electron flow called Cyclic phosphorylation, finally
oxidizing P700 and yielding ATP.
➢ Non Cyclic Phosphorylation
Excited electrons from Photosystem II (P680) moves through a series of
electron flow to Photosystem I (P700) called Non Cyclic
phosphorylation, finally oxidizing P680 and yielding
NADPH.
➢ Water photolysis
Photolysis of water contributes the electron needed to reduce
P
680.
+
H O + Light energy--------->1/2O + 2H + 2 Electrons
2 2
• Non-cyclic Electron Transport System
• The light energy of specific wavelengths is
absorbed by chlorophylls and accessory pigments
of PS-II. These pigments transfer their absorbed
energy to PS-II reaction centre - P680 (chlo 680).
This centre become photo excited and exudes an
electron with a gain of energy (23 K Cal/mol).
The electron is immediately accepted by the
primary acceptor quinone. The reaction centre
comes to ground state by getting an electron
from photo-oxidation of water. (The overall
process involving conversion of light energy into
chemical form is called quantum conversion).
• Photolysis of Water
• The PS-II reaction centre (P680) by transferring
electron to primary acceptor becomes oxidised.

• The overall process is called photo-oxidation of water.

++ + -
It requires the presence of Mn , Ca and Cl , a water
oxidising enzyme and an unknown substance Z. It is
believed that oxygen evolves as oxygen gas. Electrons
are accepted by PS-II reaction centre through unknown
+
substance Z and H temporarily stay in the thylakoid
space (loculus).The high energy electrons that leave PS-
II are captured by Q which sends them to an electron
transport system consisting of PQ, cytochrome
complex, PC. Every time electron passes from donor to
acceptor, the reduced donor id oxidised and the
acceptor is reduced. The electrons of plastocyanin are
picked by PS-I.
Z Scheme of Light Reaction
Cyclic Photophosphorylation

• The electrons released by P700 of PS-I in the presence of light are

taken up by the primary acceptor and are then passed on to
ferredoxin (Fd), plastoquinone (PQ), cytochrome complex,
plastocyanin (PC) and finally back to P700 i.e., electrons come
back to the same molecule after cyclic movement The cyclic
photophosphorylation also results in the formation of ATP
molecules just like in non - cyclic photo phosphorylation.
• As the electrons move downhill in the electron transport chain,
they lose potential energy and ATP molecules are formed in the
same way as in mitochondria during respiration.
• During cyclic photophosphorylation, electrons from photosystem - I
are not passed to NADP from the electron acceptor. Instead the
electrons are transferred back to P700. This downhill movement of
electrons from an electron acceptor to P700 results in the formation of
ATP and this is termed as cyclic photophosphorylation. It is very
important to note that oxygen and NADPH2 are not formed during cycle
photophosphorylation.
 Dark Reaction/calvin Cycle
Basic function of this cycle is reducing CO2 to carbohydrates in
all photosynthetic eukaryotes which takes place in Stroma of
Chloroplast. Also known as Photosynthetic Carbon Reduction
Cycle, or Reductive Pentose Phosphate (RPP) Cycle.
There are three basic stages of Calvin Cycle:
➢ Carboxylation
Carboxylation of the CO2 acceptor Ribulose-1,5-
bisphosphate, forms two molecules of 3 -Phosphoglycerate
(PGA), the first stable intermediate of the Calvin cycle.
➢ Reduction
Reduction of 3-PGA forms Glyceraldehyde-3-Phosphate,
a carbohydrate.
➢ Regeneration
Regeneration of the CO2 acceptor ribulose-1,5-
bisphosphate occurs from Glyceraldehyde-3-Phosphate.
Stoichiometry of the Dark Reaction
In the six turns of the Calvin Cycle, 6 CO 2
molecules have entered and bound to 6 molecules
of RuBP to yield 12 molecules of G3P.
To "fix" each CO2 molecule into G3P requires 2
ATP and 2 NADPH; an additional ATP is required
in the regeneration of RuBP.
 The Four - Carbon Pathway
C4 plants like maize, sugarcane; pearl millet, etc. have evolved
a wonderful mechanism to avoid photorespiration, which is
considered a wasteful process.
In these plants three or 2 types of photosynthetic cells namely
mesophyll cells and bundle sheath cells. The bundle sheath
cells are arranged in a wreath like manner, thus calling
them as kranz anatomy (kranz : wreath).
One more speciality of C4 plants is that they contain dimorphic
chloroplasts, that is chloroplasts in bundle sheath cells are
agranal. The presence of two types of cells allows the
occurrence of light reactions and carbon reactions separately.
In C4 plants, light reactions occur in the mesophyll cells,
whereas CO2 assimilation occurs in the bundle sheath cells.
This type of separation does not allow O2 released in
mesophyll cells to escape in to the bundle sheath cells. This
prevents the oxygenation of RuBP, which is present in the
bundle sheath cells.
➢ Transverse section of maize leaf showing the arrangement of mesophyll
and bundle-sheath cells. The C4 pathway takes place in the mesophyll cells,
and the C3 pathway (Calvin cycle) operates in the bundle-sheath cells. Both
types of cells contain chloroplasts.
To further avoid photorespiration, C4 plants have
evolved a CO2 concentrating mechanism called the C 4
pathway. The main objective of C4 pathway is to build
up a high concentration of CO2 in the vicinity of
Rubisco, thus favouring carboxytition and suppressing
photorespiration.
Many plants like maize and sugar cane, are far more
efficient at taking up CO2 than C3 plants. The CO2
acceptor in C4 plants is phosphoenolpyruvate (PEP).
PEP reacts with CO2 to form oxaloacetic acid which is
reduced by NADPH to form malic acid. The malic acid
then reacts with RUBP to form pyruvic acid and PGA.
The pyruvic acid is then phosphorylated by ATP to
regenerate PEP while PGA is converted to triose
phosphate as far as C3 plants. These reactions are
called the Hatch - Slack pathway.
These plants, however first fix carbon dioxide in the
four - carbon compound oxaloacetic acid. C4 plants
have two rings of cells surrounding the vascular
bundles. The outer mesophyll cells fix CO2 using
PEP and pass malic acid to the inner bundle sheath
cells. Here CO2 is released for acceptance by RUBP
and the eventual formation of triose phosphate. The
photosynthetic efficiency of C4 plants is due to their
ability to fix CO2 in environmental conditions where
CO2 is the limiting factor. The C4 photosynthetic
pathway is more efficient than the C3 pathway due to
the absence of photorespiration in C4 plants.
 Differences between C3 and C4 Plants

C3 C4

CO2 acceptor is 5c RUBP CO2 acceptor is 3c PEP

CO2 fixing enzyme is RUBP CO2 fixing enzyme is PEP

carboxylase carboxylase

First product of photosynthesis is First product of photosynthesis is

PGA (3c) oxaloacetic acid (4c)

CO2 fixation occurs once only CO2 fixation twice - Kranz anatomy
 Crassulacean acid metabolism (CAM)
CAM refers to a mechanism of photosynthesis that occurs only
in succulents and other plants that grow in dry conditions.
a) In CAM plants, CO2 is taken up by the leaves, which are
present on green stems through the stomata. But the stomata
remain open only during the night. During the day it remains
closed to conserve moisture.
b) The CO2 taken is fixed in the same way as in C4 plants,
to from malic acid which is stored in the vacuole.
c) This malic acid formed during the night is used as a source of
CO2 during the day for photosynthesis to proceed via the C3
pathway.
Thus CAM is a kind of adaptation in certain plants (grown in dry
conditions) to carry out photosynthesis without much loss of
water, which is otherwise unavoidable in C3 and C4
photosynthetic mechanisms.
➢ CAM pathway showing carbon dioxide uptake through open stomata during
night and its utilisation for the formation of malic acid which is stored in the
vacuole. During day, the malic acid is decarboxylated to release carbon dioxide
which is re-fixed to produce starch inside choloroplast via C3 calving cycle.
 Photorespiration
It has been observed that a high concentration of oxygen,
inhibits photosynthesis. This is due to the reason that the
CO2 fixing enzyme RUBP carboxylase not only accepts CO2
but can also combine with O2. Since the reaction is an
oxygenation reaction, the same enzyme is called RUBP
oxygenase.In the chloroplast, phosphoglycolate is unstable
and is converted to glycolate. PGA is used up in the calvin
cycle. The peroxisomes present in the cell convert the
glycolate into glyoxylate and then into glycine. Glycine is
converted to serine and CO 2 in the mitochondria. The serine
thus formed is converted to glycerate, through a series of
reactions which occurs in the peroxisome.
➢ This process is known as photorespiration or photosynthetic
carbon oxidation cycle.
➢ Photorespiration is defined as a light dependent uptake of O2 and
output of CO2.
Photorespiration involves oxygenation of ribulose-bisphosphate catalysed by the
enzyme Rubisco. The reactions lead to the formation of 2 -phosphoglycolate,
oxidation of which results in release of CO 2 . In contrast, carboxylation, also
catalysed by Rubisco, results in fixation of CO2 for the production of carbohydrate
Or
Photorespiration is also defined as the respiration initiated in the chloroplasts, in
the presence of light only.
➢ The main features are:
• The carbon lost is not retrieved. The plant may lose as much as half of the
CO2 fixed during photosynthesis by this process.
• No energy rich compound is produced and rather is a wasteful loss of ATP and
NADPH.
• With the increase in O2 concentration, affinity of RUBP carboxylase for O2
increase and for CO2 decreases. Therefore increased oxygen level
increases, photorespiration, while increased carbon dioxide level decreases
photorespiration.
• It involves three organelles - chloroplast, perioxisomes and mitochondria.
• Increase in temperature leads to more photorespiration that means more loss
of photosynthetically fixed carbon.
• It occurs in light and has no relation with the normal respiration.
• It decreases the yield of C3 plants.
 Factors Affecting Photosynthesis
Photosynthesis is influenced by two categories of factors - external or
environmental and internal or plant factors. According to the law of
limiting factors, put forward by F.F.Blackman in 1905,
photosynthesis is limited by the most limiting factor. This means
that, at any given time, only one factors is the most limiting factor
among all and this factor determines the rate of photosynthesis.
External Factors
1) Carbon dioxide
In C3 plants, the rate of photosynthesis increases with an increase. In
-1
CO2 concentration upto 500 ml.l .
In C4 plants, photosynthetic rate increases with an increase in CO2
concentration. However these plants attain saturation at around 360
-1 -1
ml.l (as compared to C3 plants which is around 500 ml.l )
-1
The current level of CO2 in the atmosphere is about 360 ml.l .
Photosynthesis could be enhanced by increasing the concentration of CO2.
Compensation Point or Threshold Value
It represents that CO2 concentration at which CO 2 fixed during
photosynthesis is equal to the volume of CO2 evolved during respiration.
2) Irradiance or Light Intensity
➢
Light
Both quality and intensity of light influence the rate of photosynthesis.
As the intensity of light increases, the rate of photosynthesis also increases. At
a very high light intensity the rate of photosynthesis decreases as
chlorophyll itself is destroyed due to high intensity. This phenomenon is
called as photooxidation as it occurs in the presence of O2.
In moderate light intensities the rate of photosynthesis is directly
proportional to irradiance.
➢ High light intensity
• Beyond saturation point, light intensity reduces photosynthesis. This effect is
called solarization. It can be caused by photo-inhibition and photo-oxidation.
➢ Photo-inhibition
Highlight intensity increases the rate of transpiration and reduction in
hydration of green cells. Stomata may close. Photo inhibition is not
influenced by O2 concentration and temperature.
➢ Photo-oxidation
Due to O2 there occurs destruction or oxidation of
photosynthetic pigments and intermediate.
➢ Wavelength of light
Wavelength of light between 400 nm and 700 nm is most effective
for photosynthesis. This light is called photosynthetically active
radiation (PAR).
Comparatively more photosynthesis occurs is red and blue regions
though others have significant net photosynthesis. Light has
maximum efficiency in red and minimum in blue region. In both
these regions light is absorbed by chlorophylls. Red light favours
more carbohydrate accumulation while blue light favours more
protein synthesis.
➢ Duration of light
If light intensities are not otherwise harmful, continuous
photosynthesis can be sustained throughout 24 hours.
3) Temperature
➢ Overall, the process of photosynthesis is sensitive to higher temperatures.
This is because the enzymes involved in photosynthesis becomes inactive at higher
temperatures. Low temperatures also inactivate the enzymes. Thus an optimum
temperature is preferred.
➢ o
The optimum temperature lies between 15 - 35 C.
o

➢ Oxygen
Small quantity of oxygen is required for optimum photosynthetic
electron transport.
➢Soil water
If water availability in the soil decreases, plants undergo water stress. Under
the condition of water stress, the rate of photosynthesis declines, as istomata
fail to open and leaf water potential decreases.
It is observed that the rate of photosynthesis is reduced with the
decrease hydration of the leaves.
➢
5) Minerals
Nutrients
Among nutrients nitrogen influences the rate of photosynthesis. Reduction in
nitrogen supply adversely affects photosynthesis, as nitrogen forms the
basic constituent of chlorophyll. In general, all essential elements also affect
the rate of photosynthesis.
6) Leaf Factors
Leaf age, leaf angle and leaf orientation also effect the rate of photosynthesis.
Mg, Fe, Cu, Mn, Cl, S and K take part in photosynthesis.
➢Air Pollutants
They decrease the rate of photosynthesis.
Examples: Ozone, hydrogen fluoride, nitrogen oxide, sulphur dioxide, dust
particles, smoke reduce light intensity and photosynthesis.
➢
Note :
The compensation point is the point at which the rate of uptake of CO2 by a plant
is equal to the rate of output of CO2. It indicates the point at which the rate of
photosynthesis and rate of respiration are exactly equal.
➢ A limiting factor is the factor, in a chemical reaction involving several
factors, which prevents the reaction rate from increasing because it is present at its
minimum level.
➢ Temperature: At low temperature the activity of RuBisCo will limit the
uptake of CO2 by RUBP. Thus the rate of formation of PGA and PGAL will be
reduced.
➢ Light Intensity: Inadequate light intensity, no electrons will flow along the
electron carrier chain and no ATP synthesis, no O2, no NADPH will occur.
➢independent
CO : CO is a reactant required in the initial chemical reactions of the light
2 2
Calvin cycle.
 Internal Factors
➢ Anatomy which favours the amount of CO2 diffusion into collenchyma
cells the intensity dilution of light and the rate of photosynthesis. Thickness of
epidermis, size structure, position and frequency of stomata, distribution of
vascular tissue and intercellular spaces, position and number of chloroplasts.
➢ Chlorophyll
Essential for photosynthesis.
➢ Carotenoids
Essential to prevent photo-oxidation which reduces photosynthesis
➢ Age
Rate of photosynthesis increases with age of the leaf till maturation.
Afterwards it begins to decrease.
➢ Demand
Decreases if meristems is removed to decrease demand for
photosynthesis. It increases at the time of increased synthetic
activity.
➢
Hormones
Cytokinin and gibberellins enhance the rate of
photosynthesis while ABA decreases.
➢ Translocation
The energy rich carbon compounds formed during
photosynthesis- called photosynthates or
photoassimilates are transported to all the organs
and tissues of the plant body. This long distance
transport of photosynthates through the phloem is
called as translocation.
The photosynthates provide energy to the non-
photosynthetic tissues through respiration. In storage
organs, photosynthates are stored in the form of
starch or as other carbohydrates
 Cellular Respiration
• All living organisms require a continuous supply of
energy for carrying out various functions. The
main source of energy for all the functions, in all
living organisms is cellular respiration.
• Cellular respiration is an energy releasing,
enzymatically controlled catabolic process, which
involves a step-wise oxidative breakdown of
organic substances inside living cell. The organic
substance i.e., glucose is oxidised inside the
mitochondria to produce energy which is liberated
in a controlled manner and which is partly stored in
the high energy bonds of ATP molecules as
biologically useful energy.
 Respiratory substrates
• The organic substances, which are catabolised in the
living cells to release energy are called as respiratory
substrates. Though any food stuff - carbohydrate, fat
or protein may act as a respiratory substrate, the
common respiratory substrate is glucose.
• Glucose is used as respiratory fuel because
a) it can be oxidised easily
b) it is present abundantly.
• Fats are used as respiratory substrates in some cases.
Proteins are used as respiratory substrates only when
carbohydrates and fat reserves have been used up.
• When proteins are used as respiratory fuels, it is
called as protoplasmic respiration.
• When fats and carbohydrates are used as
respiratory fuel, it is called as floating respiration.
 Types of Respiration
Cellular respiration is of 2 types according to its dependence
on oxygen.
• Aerobic respiration
Organic food is completely oxidised with the help of oxygen
into carbon dioxide and water. This occurs in the mitochondria.
• Anaerobic respiration
When food is oxidised without using molecular oxygen it is
called anaerobic respiration. Anaerobic respiration is exceptional
in some cases as in the tape worms living inside the human
intestine. But temporary anaerobic respiration may occur even
in our body in the fast working skeletal muscles, as in running,
swimming etc. The fatigue experience is due to lactic acid
accumulated in the muscles, due to the shortage of oxygen, a
condition called as oxygen debt. When one rests after the
exercise the lactic acid gets slowly oxidized by the oxygen
available later and then the debt is cleared.
Anaerobic respiration is also called fermentation.
 Mechnism of respiration
• Glucose is a carbohydrate - a compound of carbon
and hydrogen. The bonds between the carbon and
the hydrogen atoms are very strong. In the cells,
the substrate, often glucose, is broken down into
carbon dioxide and water in the presence of
oxygen. This process breaks the bonds between
carbon and hydrogen and thus releases energy. This
is called respiration. It has three main steps
• Glycolysis
• Kreb's Cycle
• Electron transport chain
 Glycolysis

This takes place in the cytoplasm of the cells. The glucose is converted to two molecules
of pyruvic acid. No oxygen is required for this step.
• Glucose is split into two molecules of pyruvate,
• During glycolysis, glucose, a six carbon sugar, is split
into two three-carbon sugars. These smaller sugars
are oxidized and rearranged to form two molecules of
pyruvate, the ionized form of pyruvic acid.
Each of the ten steps in glycolysis is catalyzed by a
specific enzyme.
These steps can be divided into two phases:
an energy investment phase (where the cell invests
ATP to provide activation energy by
phosphorylating glucose.)
• and an energy payoff phase (where ATP
is produced by substrate level phosphorylation
+
and NAD is reduced to NADH by electrons
released by the oxidation of glucose.)
 Kreb's Cycle
It takes place in the mitochondria. The pyruvic acid formed during glycolysis is oxidised and
forms carbon dioxide and water. This reaction also releases molecules called the coenzymes:
NADH2 and FADH2. The hydrogen atoms associated with these are the hydrogen atoms
released during the oxidation of pyruvic acid.
• After pyruvate enters themitochondrion via activetransport, it
is converted to acompound called acetyl coenzymeA or acetyl
CoA. This step isaccomplished by a multienzymecomplex.
• Acetyl Co-A drives the cycle of reactions which produces
hydrogen. If oxygen is present, pyruvate enters the
mitochondrion where enzymes of the citric acid cycle
complete the oxidation of the organic fuel to carbon
dioxide. The citric acid cycle has eight steps, each catalyzed
by a specific enzyme.
• The acetyl group of acetyl CoA joins the cycle by combining
with the compound oxaloacetate, forming citrate.
• The next seven steps decompose the citrate back to
oxaloacetate. It is the regeneration of oxaloacetate
that makes this process a cycle.
• Three CO2 molecules are released, including the one
released during the conversion of pyruvate to acetyl CoA.
 Electron transport chain
• It takes place in Inner membrane of mitochondria
• Hydrogen dives a series of reactions which release
enough energy to make ATP.
• In the electron transportchain, the electrons move from
molecule to molecule until they combine with
molecular oxygen ions to form water.
• The electron transport chain is a collection of molecules
embedded in the cristae, the folded inner membrane of
the mitochondrion.
• Electrons carried by NADH are transferred to the first
molecule in the electron transport chain, a flavoprotein.
• The electrons continue along the chain that includes
several cytochrome proteins and one lipid carrier.
• The last cytochrome of the chain, cyt a3, passes its
electrons to oxygen, which is very electronegative.