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PLANT
PHYSIOLOGY
Dr Shivani Bhargava

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SHIVANI DEMOLIVE
SHIVANI
DEMOLIVE
HISTOLOGY 101
Complete Plant Physiology
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SHIVANI DEMOLIVE
SHIVANI
DEMOLIVE
Transpiration:
Transpiration means the the loss of water in the
form of water vapours from the aerial parts of the
plants.

Transpiration is similar to evaporation in that water

from the wet surface is converted into vapours
form. It is different from evaporation in being
controlled by structural and physiological
adaptions of plants.
The minimum transpiration is found in succulent
xerophytes.

Maximum transpiration is found in mesophytes.

Types of Transpiration:
1. Stomatal transpiration : It is the transpiration
which occurs through the stomata present on
the leaves of the plants and delicate organs. About
50%-97% transpiration occurs through t he
stomata.

2. Cuticular transpiration : Water vapour are

also lost directly from the outer walls of the
epidermal cells through the cuticle. Cuticle is a
wax like layer present on epidermis. About 9% to
9.9% transpiration is cuticular.
3. Lenticular transpiration : Lenticels are
aerating pores in the cork of the woody stems,
fruits and twigs. water vapours are lost through
these openings. Amount of water vapours lost
through lenticles is usually significant. however,
approx. 0.1% to 1% is total water lost.

4. Bark transpiration : This occurs through the

bark of woody stem.
Stomata are found on the aerial delicate organs
and outer surface of the leaves in the form of
minute pores. Stomatal pore is surrounded by two
specialised epidermal cells called as guard cell.
They are kidney shaped. The number of guard
cells are two.
* The structure of guard cells in monocots
(Gramineae) is dumbel shaped
* Guard cells are epidermal cells. But due to
presence of chloroplast they are different from that of
epidermal cells.
* The outer wall of the guard cells is thin and
elastic, while inner wall is thick and elastic.
* Guard cells are surrounded by some specialized
epidermal cells called subsidiary cells or accessory
cells.
* Stomata are found on both upper and lower
surface. Stomata attached with air chambers and
forms a cavity is called sub-stomatal- cavity.
* In xerophytic plants position of stomata is deep in
the surface of the leaf. Stomata are present in this
position are called sunken stomata.
Factors affecting the rate of transpiration :-
Factors affecting the rate of transpiration are divided
into two types :-
[A] Internal Factors
[B] External factors (Environmental factor)
[A] INTERNAL FACTORS :-
These factors are concerned with structure of plants.
These are of following types :-
[1] Transpiring area :-
Pruning increase the rate of transpiration per
leaf but overall reduce the transpiration.
[2] Anatomical characteristics of leaf and leaf
orientation :
Several structures of leaf effect the transpiration
as follows :-
Stomatal characteristics :-
Transpiration is effected by the structure of stomata,
position of stomata, distance between the stomata,
number of stomata per unit area and activity of the
stomata.
[3] Water status of Leaves
[4] Root - Shoot Ratio
This is the most important factor. The rate of
transpiration is higher in low atmospheric humidity
while at higher atmospheric humidity, the atmosphere
is moistened, resulting decreasing of rate of
transpiration.

Therefore, the rate of transpiration is high during the

summer and low in rainy season.
 The value of Q10 for transpiration is 2. It means by
increasing 100C temperature, the rate of transpiration is
approximately double. (By Loftfield)
 Water vapour holding capacity of air increased at high
temperature, resulting the rate of transpiration
increased.
 On contrary vapour holding capacity of air decreased
at low temperature so that the rate of transpiration is
decreased.
[3] Light :-
 Light stimulates, transpiration by heating effect on
leaf.
 Action spectrum of transpiration is blue and
red.
 Rate of transpiration is faster in blue light than
that of red light. Because stomata are completely
opened as their full capacity in the blue light.
 Transpiration is less in constant air but if wind
velocity is high the rate of transpiration is also high,
because wind removes humid air (saturated air)
around the stomata.

 Transpiration increases in the beginning at high

wind velocity [30 - 35 km./hour] But latter on it cause
closure of stomata due to mechanical effect and
transpiration decrease.
[5] Atmospheric Pressure :-
At low atmospheric pressure, rate of the diffusion
increase which increase the rate of transpiration.
The rate of transpiration is found maximum at high
range of hills.
By carrying a plant from Kota, to hill station, rate of
transpiration increased.
Transpiration ratio (TR) :
 Moles of H2O transpired/moles of CO2
assimilated
 Ratio of the loss of water to the photosynthetic
CO2 fixation is called TR.
 TR is low for C4 plants (200-350) while high for
C3 plants (500-1000). It means C4 conserve
water with efficient photosynthesis.
 CAM plants passes minimum TR (50-100)
[6] Anti transpirants :-
Chemical substances which reduce the rate of
transpiration are known as antitranspirants. Anti
transpirants are as follows :-

 Phenyl Mercuric Acetate [PMA], Aspirin,

(Salicylic acid), Abscisic Acid [ABA], Oxi -
ethylene, Silicon oil, CO2 and low viscous wax

 PMA closed the stomata for more than two weeks

partially.
Antitranspirants are used in dry farming.
Transpiration and Photosynthesis - a
Compromise -
 Creates transpiration pull for absorption and
transport of plants.
 Supplies water for photosynthesis.
 Transports minerals from the soil to all parts of
the plant.
 Cools leaf surfaces, sometimes 10 to 15
degrees, by evaporative cooling.
* How do plants absorb water
forms of water

* Water is mainly obtained from the rain. Some of

the water goes into the water reservoirs. This is
called run off water Res of the water present in the
soil is of following types
Gravitational water : Form of water which reaches
in earth up to water table due to he gravitational
force after rainfall. This form is not available for
plants naturally but can be made available by
mechanical methods or by tube well irrigation.
Exceptionally some plants can absorb this water
Calotropis, Prosodies, Capris etc.
Hygroscopic water :
A thin film of water is tightly held each soil particle
called as hygroscopic water. This water is also not
available to the plants.
Chemically combined water :
The amount of water present with some chemical
compounds, which are present in the particles of
soil is called chemically combined water. This is also
not available to the plants,
Capillary water :
Water exists between soil particles in small capillary
porex, it is called capillary water. Plants absorb only
this form of water.
Holard: It is the total amount of water present in the
soil
Chresard : This is the water available to the plants.
Echard : This is the water not available to the plants
The absorption of water and minerals takes place
through root hairs. The root hairs increase the
absorption area of root.
* Once water is absorbed by root hairs, It can
move deeper into root layers by two distinct
pathways
(a) Symplast :
* This is the living path, System of interconnected
protoplasts through plasmodesmata.
* During sypmpastic movement . the water travels
through the cells, their cytoplasm and
plasmodesmata (during intercellular movement).
Symplastic movement may be aided by
Cytoplasmic streaming.
* Water has to enter the cells through the cell
membrane, Hence the movement is relatively
slower. It is also called trans membrane pathway.
(b) Apoplast :-
* This is the non living path in plants. Cell wall
intercellular space and xylem cavity associate
together to form applets.
* Flow of water from root hair to cortex is both
apoplectic and simplistic. In Endodermis there are
caesarian strips so further flow of water occurs only
by Simplest.
* Apoplast does not provide any barrier to water
movement so it is relatively faster than the
syrsplast and in apoplast. water movement is
through mass flow.
* Most of the water flow in the roots occurs via
the apoplast since the cortical cells are loosely
packed and hence offer no resistance to water
movement.
Long Distance Transport ( Translocation)
• Long distance transport of water, minerals and
food occurs generally by mass or bulk flow
system.

* Mass flow is the movement of substances in bulk

or en masse from one point to another as a result of
pressure difference between the two points.
* It is a characteristic of mass flow that substances,
whether insolution or in suspension , are swept along at
the same pace, as in a flowing river. This is unlike
diffusion where different substances move
independently depending on their concentration
gradients.
* Bulk flow can be achieved either through a positive
hydrostatic pressure gradient (pushing pressure) or a
negative hydrostatic pressure gradient ( pulling
pressure)
Water Movement up A plant (Ascent of Sap)
* Upward movement of absorbed water against
the gravitational force upto parts of plants is called
as ascent of sap. (Translocation of sap water +
minerals) by xylem
* Experiment of Balsam plant by using eosindye
prove that xylem is water conducting tissue of the
plants.
Mechanism of ascent of sap :
Cohesion - tension- transpiration pull model : By
Dixon & Jolly.

* Most- accepted or universally accepted theory

for explaining mechanism of ascent of sap.
* According to it following components are
involved in ascent of sap
* Cohesion : Mutual attraction between the water
molecules is known as cohesion. which helps to
forms a continuous water column in xylem
elements.

* Adhesion : Attraction between xylem walls &

water molecules is called adhesion force, which
helps in upward maintenance of water column in
the xylem.
Surface tension
Transpiration pull : A negative pressure (pulling
pressure) develops in xylem due to rapid transpiration in
leaves, this is called transpiration pull which is
responsible for the pulling of water column in xylem.
* Ascent of sap is constitutive effect of cohesion,
surface tension, adhesion & transpiration pull.
* Measurements reveal that the force generated
by transpiration can create pressure sufficient to
lift a xylem sized column of water over 130 meters
high. (with the rate of about 15 meters per hour).
* Cohesion, Adhesion and surface tension
properties give water high tensile strength means
ability to resist pulling force and also give water
high capillarity means ability to rise in thin tubes, In
plants capillarity is aided by the small diameter of
the tracheids and vessels.
Root Pressure :-
* As various ions from the soil are actively
transported into the vascular tissues of the roots,
water follows(its potential gradient) and increases
the pressure inside the xylem . This positive pressure
is called root pressure , and can be respondible for
pushing up water to small heights in the stem.
* The greatest contribution of root pressure may
be to re-establish the continuous chains of water
olecules in the xylem which often break under the
enormous tensions created by transpiration.
* Root pressure only provides a modest push in
the overall process of water transport.
* Root pressure does not account for the majority
of water transport: most plants meet their need by
transpiratory pull.
PHLOEM TRANSPORT (FOOD
TRANSLOCATION IN PLANTS)
* Food/ organic material conduction in plants
occurs by phloem. (Proved by girdling experiment)
* Generally green photosynthetic plant parts aci
as source like leaves while non photosynthetic
parts like root , shoot, fruits act as sink. Food
conduction occurs from source to sink.
* Food transfer depends on requirement of
plant and seasonal activities for example in
germinating potato tuber, tuber acts as source and
developing buds acts as sink , similarly in early
spring roots act as source and developing buds as
sink
* Translocation of food occurs in the form of
sucrose because it is a non- reducing sugar and
chemically inert in it s pathway of conduction.
* Phloem sa is mainly water and sucrose. But
other sugars, hormones and amino acids are also
transported or translocate through phloem.
* Pressure flow/ mass flow hypothesis of food /
sucrose translocation

* Given by E. Munch (1930) , this is the most

accepted theory of food conduction in plants.
Uptake and Transport of Mineral Nutrients
Plants obtain their carbon and most of their oxygen
from CO2 in the atmosphere. However, their remaining
nutritional requirements are obtained from minerals
and water for hydrogen in the soil.
Uptake of Mineral lons
Unlike water, all minerals cannot be passively absorbed
by the roots. Two factors account for this:

(i) Minerals are present in the soil as charged particles

(ions) which cannot move across cell membrances and

(ii) The concentration of minerals in the soil is usually

lower than the concentration of minerals in the root.
Therefore, most minerals must enter the root by
active absorption into the cytoplasm of epidermal
cells. This needs energy in the form of ATP.

The active uptake of ions in partly responsible for

the water potential gradient in roots, and therefore
for the uptake of water by osmosis.

Some ions also move into the epidermal cells

passively.
lons are absorbed from the soil by both passive and
active transport. Specific proteins in the
membranes of root hair cells actively pump ions
from the soil into the cytoplasms of the epidermal
cells.

Like all cells, the endodermal cells have many

transport proteins embedded in their plasms
membrane; they let some solutes cross the
membrane, but not others.
Transport proteins of endodermal cells are control
points, where a plant adjusts the quantity and types
of solutes that reach the xylem.

Note that the root endodermis because of the layer

of suberin has ablility to actively transport ions in
one direction only.
 Translocation of Mineral Ions
After the ions have reached xylem through active or
passive uptake. or a combination of the two, their further
transport up the stem to all parts of the plant is through
the transpiration stream.

The chief sinks for the mineral elements are the growing
reglons of the plant, such as the apical and lateral
meristems, young leaves, developing flowers, fruits and
seed, and the storage organs. Unloading of mineral ions
occurs at the fine vein endings through diffusion and
active uptake by these cells.
 Mineral ions are frequently remobilised, particularly
from older senescing parts. Older dying leaves export
much of their mineral content to younger leaves.

 Similarly before leaf fall in decidous plants, minerals

are removed to other parts.

 Elements most readily mobilised are phoshorus,

sulphur, nitrogen and potassium. Some elements that
are structural components like calcium are not
remobilised.
An analysis of the xylem exudate shows that
though some of the nitrogen trabels as the xylem
exudates shows that though some of the form as
amino acids and related compounds. Similarly,
small amounts of P and S are carried as organic
compounds.

In addition, small amount of exchange of

materials does take place between xylem and
phloem.
Hence, it is not that we can clearly make a
distinction and say categorically that xylem
transports only inorganic nutrients while phloem
transports only organic materials, as was
traditionally believed.
PHLOEM TRANSPORT : FLOW FROM SOURCE
TO SINK

Food, primarily sucrose, is transported by the

vascular tissue phloem from a source to a sink.
Usually the source is understood to be that part of
the plant which synthesises the food, i.e., the leaf,
and sink, the part that needs or stores the food.
.
But, the source and sink may be reversd
depending on the season, or the plant's needs.
Sugar stored in roots may be mobilised to
become a source of food in the early spring when
the buds of trees, act as sink ; they need energy
for growth and development of the photosynthetic
apparatus.

Since the source-sink relationship is variable, the

direction of movement in the phloem can be
upwards or downwards, i.e., bi-directional.
This contrasts with that of the xylem where the
movement is always unidirectional, i.e., upwards.
Hence, Unlike one-way flow of water in
transpiration, food in phloem sap can be
transported in any required direction so long as
there is source of sugar and a sink able to use,
store or remove the sugar.

Phloem sap is mainly water and sucrose, but other

sugars, hormones and amino acids are also
transported or translocated through phloem
The Pressure Flow or Mass Flow Hypothesis

 The accepted mechanism used for the

translocation of sugars from source to sink is
called the pressure flow hypothesis.
 As glucose is prepared at the source (by
photosynthesis) it is converted to sucrose
(dissacharide).
 The sugar is then moved in the form of sucrose
into the companion cells and then into the living
phloem sieve tube cells by active transport.
This process of loading at the source produces a
hypertomic condition in the phloem.

 Water in the adjacent xylem moves into the

phloem sap will move to areas of lower pressure. At
the sink osmotic pressure must be reduced.

 Again active transport is necessary to move the

sucrose out of the phlowm sap and into the cells
which will use the sugar-converting it into energy,
starch, or cellulose.
As sugars are removed, the osmotic pressure
decreases and water moves out of the phloem.

To summarise, the movement of sugars in the

phloem begins at the source, where sugars are
loaded (actively transported) into a sieve tube.
Loading of the phloem sets up a water potential
gradient that facilitates the mass movement in the
phloem.
Phloem tissue is composed of sieve tube cells.
which from long columns with holes in their end
walls called sieve plates.

Cytoplasmic strands pass through the holes in the

sieve plates, so forming continuous filaments.

As hydrostatic pressure in the phloem. Meanwhile,

at the sink, incoming sugars are actively tranported
out of the phloem and removed an complex
carbohydrates.
The loss of solute produces a high water potential
in the phloem. and water passes out, returning
eventually to xytem.

A simple experiment called girdling, was trunk of a

tree a ring of bark up to a depth of the phloem
layer, can be carefully removed.
In the absence of downward movement of food the
protion of the bark above the ring on the stem
becomes swollen after a few weeks.

This simple experiment shwos that phloem is the

tissue responsible for translocation of food ; and
that transport takes place in one direction, i.e.,
towards the roots.
GUTTATION
 The term "Guttation" was coined by
Burgerstein.
 Loss of water from the uninjured part or
margin of leaves of the plant in the form of water
droplets is called as guttation.

 The process of guttation take place due to root

pressure, develop in cortex cells of root.
Generally guttation occurs during mid night or early
morning.
Parenchymatous and loose tissue are lie beneath
the hydathode, which are known as epithem or
transfer tissues.

Exuded liquid of guttation along with water contains

some organic and inorganic (dissolved)
substances. It means it is not pure water.

Guttation occurs from the margins of the leaves

through the special pore (always open) like structure
are called hydathodes or water stomata.
Normally, guttation process is found in herbacious
plants like grasses, tomatos, balsum,
Naustertium, Colocasia, Sexifraga and in some
of the plants of Cucurbitaceae family.
Bleeding
Fast flowing of liquid from the injured or cut
parts of the plants is called bleeding or
exudation.
This process takes place due to high root pressure.
Sugar is obtained from the Sugar mapple by this
process.
The highest bleeding is found in Caryota urens
(Toddy palm) (about 50 liter per day).
Bleeding is important in economic biology, because
Opium, Latex of rubber is obtained by this.
Photosynthesis
· Stephen Hales (1727):- Recognised the
importance of air (CO2) and light for
photosynthesis (nourishment) in plants. He is
considered as discoverer of photosynthesis and
"Father of plant physiology"
(Father of Indian plant physiology– J.C. Bose)
· J. Priestley (1770):- He carried out very
interesting exp.with Bell jar. Rat. Pudina & Candle.
He came to conclusion that purify the air
(gaseous exchange). during photosynthesis.
Discoverer of oxygen. Phlogision Þ Bad air from
candles, Dephlogistion Þ Good air
Experiment by Joseph Priestley :
· 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 observed that a candle burning in a closed
space - a bell jar , soon gets extinguished. Similarly ,
a mouse would soon suffocate in a closed space. he
concluded that a burning candle or an animal that
breather the air, both somehow, damage the air, But
when he placed a mint plant in the same bell jar, he
found that the mouse stayed alive and the candle
continued to burn.
· Prisetley hypothesised as follows: Plants
restore to the air whatever breathing animals and
burning candles remove.
· Jan Ingenhousz (1779):- Importance of light
and green colour. O2 is released in the presence
of light by green parts.

Experiment by Jan Ingenhousz :

· Using a similar setup as the one used by
Priestley, but Priestley , but by placing it once in
the dark and once in the sunlight. Jan
Ingenhousz (1930-1799) showed that sunlight is
essential to the plant process that somehow
purifies the air fould by burning candles or
breathing animals.
· Lngenhousz 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.

· J.V. Sachs (1854) : Recognised the relation

among photosynthesis, chloroplast and starch.
First visible product of photosynthesis is starch.
· Engelmann (1888)- Described Action spectrum
of photosynthesis by using Cladophora, Spirogyra
algae and aerobic bacteria experiment.
Experiment by Engelmann:
· T.W. Engelmann (1843-1909) Used a prism to
split light into its spectral components and then
illuminated a green algae. Cladophora, placed in a
suspension of aerobic bacteria. The bacteria were
used to detect the sites of O2 evolution. He
observed that the bacteria accumulated mainly in
the region of red and blue light of the split
spectrum. A first action spectrum of
photosynthesis was thus described. It resembles
roughly the absorption spectrum of chlorophyll a
and b.
· Van Neil (1931): – O2 release from water not
from CO2. Experiment on purple and green
sulphur bacteria.
H2A = Suitable oxidisable compound or H-
Donor
· Ruben, Hassid & Kamen (1941) :-Used O18
(radioisotopic technique) to show experimentally
that O2 is photosynthesis released from water.
· Robert Hill & Bendal (1937) : - Detailed study of
light reaction in isolated chloroplast of Stellaria
plant.

· Photolysis of H2O is the chief role of chloroplast

and evolution of O2 occurs only in the presence of
suitable e– acceptor, from water in photosynthesis.
(Hill-reaction).Electron acceptors = Potassium
Ferricyanide, NADP+ and DCPIP (Dichlorophenol
indophenol - a dye)
"Photosynthesis is a photo–biochemical process
(anabolic & endergonic) in which organic
compounds (carbohydrates) are synthesised from the
inorganic raw material (H2O & CO2) in presence of
light & pigments. O2 is evolved as a by product".
Light energy is conserved into chemical energy
by photosynthesis.
* 90% of total photosynthesis is carried out by
aquatic plants (85% algae) & 10% by land plants.
* First true & oxygenic photosynthesis started in
cyanobacteria. (BGA)
* In the Cuscuta & fungi photosynthesis is absent.
Euglena is photosynthetic organism & is link
between animal & plants.
* Roots of Tinospora & Trapa are assimilatory or
photosynthetic.
* Absorption spectrum of photosynthesis is blue &
red light. (maximum absorbed part of spectrum)
* Action spectrum of photosynthesis is red & blue
light. (most effective in reaction)
* Rate of photosynthesis is higher in red wavelength
of light, but highest in white light (Full spectrum),
than monochromatic light.
Important scientific contribution :
* According to Van Niel, oxygen comes from water
in photosynthesis.

Ruben, Hassid and Kamen (1941) :– Used O18 to

show experimentally that O2 in photosynthesis
comes from water.
Existence of two steps in Photosynthesis –
* Blackman discovered dark reaction (By study of
Q10 value or tempereture coefficient).

* Calvin and Benson gave cyclic pathway for this,

thus dark reaction is called as Calvin cycle OR
C3–cycle.

* Q10 (Tempereture coefficient) for light reaction is

one, while Q10 for dark reaction is between
2-3. (By Vont Hoff).
* Q10 means the doubling of rate of reaction, which
involves chemicals, on 10°C rise in temperature in
it's optimum range.
* Experimental evidences for blackman findings
were given by Warburg 1920. He carried out
intermittent light experiment on Chlorella. (by
using flash light)
* The product of photosynthesis has been found
greater in intermittent light (i.e., light given after
intervals of dark periods) than in continuous light.)
* This is due to the fact that light reactions are
faster than the dark reaction.

In continuous light product of light reactions

(ATP and NADPH2) are not consumed at the
same rate as in subsequent dark reaction. Thus
dark reaction is rate limiting step of
photosynthesis.
* Photosynthesis – (i) Light reactn/Hill reactn
(ii) Dark reaction/Blackman reactn.
* Hill Reaction – Experiment on isolated
chloroplast (Stelaria plant) study of light reaction,
which is called as Hill Reaction.
* O2 gas liberated from photolysis of H2O, only in
the presence of suitable e– acceptor. (DCPIP
(Dichlorophenol Indophenol- a dye), ferricyanide,
NADP+ – Hill reagents
* Emerson & Arnold – worked on Chlorella and gave the
concept of two photosystem or two pigment systems.
When they gave only monochromatic light, longer than
680 nm wavelength, then quantum yield is suddenly
dropped down, this event is called as red drop.
* When Emerson gave light, shorter and greater than 680
nm (combined light) then photosynthetic, activity increases,
this is called as Emerson effect or enhancement effect.
(i) 680 nm­r PS – I (cyclic process) red drop
appears.
(ii) 680 nm¯ + 680­nm (Mixed light) rBoth cyclic & non
cyclic operates. (Emerson effect)
Quantum requirement –
The number of light Quanta or photons required for
the evolution of 1 mol. of O2 in photosynthesis = 8
Quantum Yield –
The number of oxygen molecule evolved by one
quantum of light in photosynthesis is called as
Quantum yield. Emerson calculated that the
quantum requirement is 8. Hence the quantum yield
is 0.125 or 12.5%)
* Many pigment present in photosynthetic cells. PSU
(Photosynthetic units) presents on thylakoid
membranes, are made up of 230-400 molecules of
various pigments, called Quantasomes by Park &
Biggins.
* The PS II is located in the appressed region of
granal thylakoids and PS I in non appressed region
of grana and stroma thylakoids.
* PS I located - on both granum & intergranum
(Stroma thylakoid), (P-700, 680 nm­, Cyclic ETS).
* PS II located - on only granum, (P-680, 680 nm¯,
non cyclic ETS).
* Chlorophyll – a C55H72O5N4Mg - CH3 grp. at IIIrd
C of IInd pyrrole ring.
* Chlorophyll – b C55H70O6N4Mg - CHO group at IIIrd
C of IInd pyrrole.
* Chlorophylls are magnesium porphyrin compounds.
Porphyrin ring consists of four–pyrrole rings
(Tetrapyrrole).
* Chlorophyll molecule has a Mg–porphyrin head
and alcoholic phytol tail. Head is hydrophilic and
phytol tail is lipophilic (hydrophobic).
* Phytol tail is alcoholic with one double bond.
Phytol part embeded in lipid layer.
* Chl–a and carotenes are universal pigment,
which are found in all O2 liberating cells.
* Chlorophylls are soluble only in organic solvents like
ketons, ethers etc.
- Stroma lamellae/stroma thylakoids lack PS II and
enzyme NADP reductase .
- In paper chromatography/chromatogram –
Chlorophyll 'a' appears bright or blue-green.
Chlorophyll 'b' as — yellow green/grass green.
Xanthophyll as — yellow
Carotenoids as — Yellow to yellow - orange
Mechanism of Photosynthesis
[A] Light reaction/Hill reaction/Photochemical reaction/
Generation of assimilatory powers (NADPH 2 +
ATPs)/photophase.
* Antenna or accessory pigments receive radient energy
and transfer it among themselves. This transfer of energy is
known as resonance transfer. Then antenna molecules
excited and transfer their energy to the chlorophyll 'a'
molecules of reaction centre.
It is known as inductive resonance. finally chl. 'a' of
leaf center molecules converts the light energy into
electrical energy by bringing about electric charge
separation.
(I) Cyclic ETS and Photophosphorylation
* In cyclic ETS, only PS–I (LHC–I) works, which
consists of Chl–'a' 670, Chl–a–683, Chl–'a'–695,
carotenoids, some molecules of chl–'b' & reaction
centre–Chl–'a'–700/P–700.
* Cyclic ETS OR PS–I is activated by wavelength of
light greater than 680 nm.
* It occurs at grana thylakoids and stroma thylakoids.
* During Cyclic ETS the electron ejected from reaction
centre of PS-I, returns back to its reaction centre.
* In cyclic ETS, no oxygen evolution occurs, because
photolysis of water is absent.
* NADPH2 (reducing power) is not formed in cyclic
process.
* Plastocyanin (PC) is Cu–containing blue
coloured protein in cyclic ETS.

* According to modern researches, first e– acceptor

is FRS (Ferredoxin Reducing Substance), which is
a Fe-S containing Protein. Earlier fd (Ferredoxin)
was considerd as first e– acceptor.
 (II) Z-Scheme/Non-cyclic ETS and
Photophosphorylation-
* Both PS–I and PS–II involved in non cyclic ETS.
* PS–II (P–680) consists of Chl–a–660, Chl–a–673,
Chl–a–680, Chl–a–690, Chl–b, or Chl–c or Chl–d,
carotenoids & phycobilins. Phycobilins present only
in PS II
* It occurs at grana thylakoids only.
* The e– ejected from PS–II never back to chl–a–
680 (reaction centre) & finally gained by NADP.
thus gap of e– in PS–II is filled by photolysis of
water as a result, oxygen evolution occurs in Z–
scheme.
* During noncyclic ETS produces approx 3 ATP
and 2NADPH2 (4 mol. of water is photolysed and 1
O2 released)
* 12 NADPH2 + 18 ATP are required as
assimilatory power to produce one molecule of
Glucose in dark reaction, thus 6 turns of Z–
scheme are necessary for the production of one
glucose molecule by calvin cycle.

* Additional ATP comes from cyclic ETS. (over all

54 ATP equivalents)
* Primary e– acceptor in non–cyclic reaction is PQ
or plastoquinone. Recently pheophytin (structure
like chl. a without Mg) is considered as Ist e–
acceptor in Z–scheme.
* Plastocyanin (PC) is link between PS–I and PS–
II in non cyclic ETS. (Some scientists–cyto-f)
* Final e– acceptor in Z–scheme is NADP+ (Hill
reagent)
* During Non-Cyclic ETS energy flow takes place
from PS II to PS I.
Chemiosmosis Hypothesis -
Proposed by Peter Mitchell (1961) to explain the
mechanism of ATP formation (Phosphorylation) in
chloroplast (During photosynthesis) and in
mitochondria (During respiration).

According to they hypothesis ATP synthesis is linked

to development of a proton gradient (Difference of
proton (H+) concentration) across a membrane. In
chloroplast these are membranes of the thylakoid.
The steps that cause a protein gradient to develop
(a) Since splitting of the water molecule takes place
on the inner side of the membrane the hydrogen
ions the are produced by the splitting of water
accumulate within the lumen of the thylakoids.
(b) As electron move through the photo stems , protons are
transported across the membrane. This happens
because the primary accepter of electron (Pheophytin)
which is located towards the outer side of the membrane
transfers its electron not to an electron carrier but to an H
carrier (Plastoquinone). Hence this molecule removes a
proton from the stroma while transporting an electron.
When plastoquinone passes on its electron to the
carrier (FeS protein or Cytochrome) on the inner side of
the membrane , the proton is released into the inner side
of the membrane or the lumen side of the membrane.
(C) The NADP reductase enzyme is located on the stroma
side of the membrane. Along with electrons that come
from the accepter of electrons ( Fd) of PS . Protons are
necessary for the reduction of NADP+ + H+. These
protons are also removed from the stroma.
Hence, within the chloroplast, protons in the stroma
decrease innumber ,while in the lumen there is
accumulation of protons. This creates a proton gradient
across the thylakoid membrane as well as a measurable
decrease in pH in the lumen
This gradient is important because it is breakdown of this
gradient that leads to release of energy . The gradient is
broken down due to the movement of proteins across the
membrane to the stroma through the transmembrane
channel of F0 of the ATPase
The ATPase enzyme consists of two parts: one called the F 0
is embedded in the membrane and forms a trans membrane
channel that carres out facilitated diffusion of protons across
the membrane. The other portion is called F1 and protruded
on the outer surface of the ehylakoid membrane on the side
that faces the stroma.
The break down of the gradient provides enough energy to
cause a conformational change in the F1 particle of the
ATPase which makes the enzyme , synthesise several
molecules of energy- packed ATP.

Chemiosmosis require a membrane , a proton pump/ H-

carrier, a proton gradient and ATPase. Energy is used to
pump protons across a membrane , to create a gradient or a
high concentration of protons . within the thylakoid lumen
[B] Dark Reaction/Blackman Reaction/Calvin
cycle/C3–Cycle/Biochemical phase/Carbon
assimilation/photosynthetic carbon reduction
cycle (PCR-Cycle)/Reductive pentose
phosphates pathway
* Blackman reaction is called as dark reaction,
because no direct light is required for this. Calvin
presented these reactions in cyclic manner &
thus called as Calvin cycle.
* Ist stable compound of Calvin cycle is 3C–PGA
(Phosphoglyceric acid) thus Calvin cycle is called as
C3–cycle. (First compound is unstable, 6C keto
acid)
* Study by Calvin was on Chlorella &
Scenedesmus. During his experiment he used
chromatography & radioisotopy (C14) techniques
for detecting reactions of C3–cycle.
* Rubisco (Ribulose bis-phosphate carboxylase-
oxygenase) is main enzyme in C3–cycle, which is
present in stroma & it makes 16% protein of
chloroplast. Rubisco is most abundant enzyme.
Diversity in Dark Reactions
CO2 concentrating mechanism/Co-operative
photo- synthesis/ Dicarboxylic acid cycle (DCA
cycle) /C4 cycle/Hatch & Slack Pathway
* Kortschak and Hartt first observed that 4C, OAA
(Oxaloacetic Acid) is formed during dark reaction in
sugarcane leaves.
* Hatch & Slack (Australia) (1967). Studied in
detail and proposed pathway for dark reactions in
sugarcane & maize leaves.

* First stable product of this reaction is OAA.

Which is 4C, DCA (Dicarboxylic Acid), thus Hatch &
Slack pathway is called as C4 cycle or DCA
cycle.
* C4–cycle occurs in 1500 sps. of 19 families of
angiosperm, but most of the plants are monocots,
which belong to Graminae & Cyperaceae
(Sugarcane, Maize, Sorghum, Chloris,
Sedges, Bajra, Panicum, Alloteropsis etc.) Rice
sps.
* Atriplex hastata & A. patula are temperate
sps,. which are C3–plants.
* Dicots with C4–cycle are Euphorbia sps.,
Amaranthus, Chenopodium, Boerhavia,
Atriplex rosea, Portulaca, Tribulus.
* Wheat and barley (monocot) are C3 species. rice
sp. devlopes as C4 plants by plant breeding
scientists.
* Kranz (Wreath) anatomy – Present in leaves of
C4 plants.
(i) Green bundle sheath cells (BS cells) present
around the vascular bundles.
(ii) Dimorphic chloroplasts present in leaf cells.
Chloroplast of B.S. cells or Kranz cells are larger
and without grana. Mesophyll chloroplast are
small and with grana.
* Rubisco present in BS cells, while PEPCase in
mesophyll cells.
* In the C4–Plant, C3–cycle occurs in bundle sheath
cells, while C4–cycle occurs in mesophylls.
* Thus operation of Hatch and Slack pathway
require cooperation of both photosynthetic cell i.e.
mesophyll cells and BS cells.
* Photosynthetically C4 plants are more efficient as
there is no warburg effect or photorespiration,
Because at the site of Rubisco (BS cells) no O 2 is
release & (mesophyll cells pumps more CO 2 for C3
* C4–plants found in tropical habitats and adapted
themselves, with high temperature, low water
availability and intense light.
* If concentration of O2 increases artificially, then
photorespiration may be started in C4 plants.
* First carboxylation in C4–cycle occurs by PEPCase
in mesophyll cytoplsam, while second carboxylation
or final CO2 fixation by C3 cycle occurs in bundle
sheath cells.
* Primary CO2 acceptor in C4 mesophyll is PEP
(Phosphoenol Pyruvate). (3C–compound), while
RuBp in bundle sheath cells.
* 12 NADPH2 & 30 ATP needed for production of 1
Hexose (Glucose) in C4–plants.
* Pyruvate phosphate dikinase (PPDK) (ATP ®
AMP) is a temperature sensitive enzyme of C 4
and CAM plants due to this C4 plants better
photosynthesizes at high temperature.
* C4 plant evolve by Anatomical, physiological &
Special features of C4 plants :-
(1) C4 plants are more efficient plants at present CO2
concentration.
(2) Present level of atmospheric CO2 is generally not
limiting factor for C4 plants.
(3) C4 plants posses low CO2 compensation points. (8-
10 ppm)
(4) The productivity (fertility) in C4 plants, does not
increase when CO2 concentration is increases.
because :
(a) Mesophyll cells pump more CO2 for Calvin cycle.

(b) Thus concentration of CO2 is high around the site

of Rubisco in C4 plants, thus little or no chance
of photorespiration.
CAM–Plants / Crassulacean acid metabolism /
Dark CO2 fixation / Dark acidification

* Oleary and Rouhani discovered CAM–process in

members of Crassulaceae family. Succulent
xerophytie plants.
Eg. are .– Kalanchoe, Bryophyllum, Sedum,
Kleinia, Opuntia, Crassula, Agave, Aloe,
Euphorbiasps, Pineapple, Welwitschia
(Gymnosperm) etc.
* Primary acceptor of CO2 is PEP (Phosphoenol
pyruvate) and oxaloacetic acid is the first product
of carboxylation reaction.
* In CAM plants stomata are of scotoactive type,
so initial CO2 fixation is found in night but light
reactions operates at day time. Final CO 2 fixation
(C3 cycle) occurs in day time. PEPcase induces
carboxylation reaction in night.
* PEP carboxylase & Rubisco present in
mesophyll cells. (No Kranz–anatomy)
* In CAM plants 30 ATP and 12 NADPH 2 are
required as assimilatory power for 1 glucose
synthesis.
* CAM plants exhibits ecophysiological
adaptation with xeric habits.
Photosynthetic carbon oxidation (PCO) cycle/C2 Cycle/
Photorespiration/ Glycolate-Metabolism
* The light dependent uptake of O2 & release of CO2
in C3photosynthetic cells is called Photorespiration
* RuBisCO is characterised by the fact that its active
site can bind to bota 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. (Ussually RuBiCO has a much greater
affinity for CO2 than for O2).
• Internal CO2 conc. Is constant arqund Rubis CO, but
internal O2 conc. is variable due to variable rate of
light reaction. When O2 conc. Is higher than CO2
conc. Then RubisCO perform oxygenation of RuBP,
which leads to phenomenon of photorespiration
• The reason of photorespiration is same site of
O2 evolution/ photolysis of H2O and catalytic activity
of Rubis CO, this site is chloroplast of Mesophyll
cells.
• Conditions for photorespiration- High O2, Low
CO2, High light intensity and high temperature.
* In C3 plants some O2 does bind to RuBisCO and
hence CO2 fixation is decreased. Here the RuBP instead
of being converted to 2 molecules of PGA. Binds with
O2 to form one molecule of phosphoglycerate (3C) and
one molecule of phosphoglycolate (2C).
• In the photorepirathory pathway there is neither
synthesis of sugars , nor of ATP and NADPH 2. Rather
it results in the release of CO2 with the utilisation of
ATP. Therefore, photorespiratioh is wasteful process.

• It occurs in chloroplast, peroxisome & mitochondria

* In C4 plants photorespiration does not occur. This is
because they have a mechanism that increases the
concentration of CO2 at the 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 true, this
ensures that the RuBisCO functions as a carboxylase
minimising the oxygenase activity.
* Also, in C4 plants , site of O2 evolution (mesophyll
cell) and site of RuBisCO activity(bundle sheath cell) are
different
Factors Affecting Photosynthesis
(1) Light
(a) Light Quality or wavelength ¾® Maxm
photosynthesis takes place in red light than in Blue
light. But rate of photosynthesis is highest in white
light. Minimum in green light.
(b) Light Intensity – Rate of photosynthesis is
greater in intense light than diffused light. But at
higher light intensity photooxidation
(solarization) occurs and photosynthetic
apparatus may get destroyed.
* P/R (Photosynthesis : Respiration) Ratio at mid
day is 10 : 1; but can reach upto 20 : 1 ratio. At the
time of evening & morning rate of photosynthesis
equals to respiration, this situation called as light
compensation point.
* Intensity of light, at which rate of photosynthesis,
becomes equal (or compensate) with the rate of
respiration in plants is known as light
compensation point. (Net photosynthesis or net
primary productivity at this point is zero.)
* Plants which are adapted to grow in high intensity
of light is called heliophytes & plants which are
adapted to grow in shade is sciophytes.
(c) Duration of Light – On the basis of effect of
light on plants may be LDP & SDP.
* Product of photosynthesis is greater in
intermittent light than continuous light – Warburg.
(2) Temperature –
* Optimum temp. for photosynthesis is 20–35°C
* At high temp. rate of photosynthesis decreases
due to denaturation of enzymes.
* Conifers & lichens can perform photosynthesis at
– 35°C,while thermal algae Oscillatoria at
70–80°C..
* Generally different habitat plants show, different
response to photosynthesis on a given temperature.
(3) CO2 (0.03%/314 ppm)
* An increase in CO2 concn. upto 1% rate of
photosynthesis is increased. Higher CO 2
concentration. is toxic to plant & also closes
stomata.
* C4–Plants can photosynthesize at low CO 2 concn
(upto 10 ppm). “CO2 concn at which CO2 fixation in
photosynthesis is equal to volume of CO 2 released
in respiration is "CO2 compensation point”, when
plant saturated with full light.
* CO2 compensation point for C4 plants is 8-10
(4) O2 –
High O2 concn. reduces photosynthesis due to
photorespiration.
(5) Water –
Less availability of water reduces the rate of
photosynthesis (stomata get closed)
(6) Chlorophyll–
The amount of CO2 in grams absorbed by 1 gm.
of chlorophyll in 1 hour is called as
photosynthetic number or assimilatory number
(Willstatter & Stoll).
(7) Product –
Rate of photosynthesis decreases, when sugar
accumulates in mesophyll cells.
(8) Leaf –
Various leaf factors like leaf age and leaf
orientation effect the rate of photosynthesis.
* In young & mature leaves photosynthesis is more
than old (senescent) leaves.
(9) Inhibitors –
DCMU (Diuron/Dichlorophenyl Dimethyl
Urea)CMU (Monuron), PAN, Atrazine,
Simazime, Bromocil, Isocil– inhibit the
photosynthesis by blocking PS–II. They stop e–
flow between P-680 & PQ.
* In cyclic ETS diquat, paraquat (Viologen dyes)
inhibit e– flow between P-700 & Fd.
* All these chemicals are used as herbicides,
which mostly block ETS.
(10) Minerals :- Mg and Nitrogen are essential for
structure of chlorophyll and enzymes. Thus
reduction in N2 and Mg supply to plants effects
adversely the rate of photosynthesis.
* Rubisco alone accounts for more than half of total
leaf nitrogen.
Generally all essential element affect the rate of
photosynthesis.
* Concept of three cardinal points (Von Sachs) :-
The effect of the various external factors on the
rate of biological processes  were centred around
the attempts to establish minimum, optimum and
maximum values known as cardinal points.
* Law of minimum (Liebig) :- According to it, when
a process is governed by a number of separate
factors, then the rate of process is controlled by that
factor present in minimum amount.
* Law of limiting factors – (Blackman) :- It is the
modification of Law of minimum by Liebig.
"When a process is conditioned to its rapidity by a
number of factors, then rate of process is limited
by the pace of the slowest factor" (CO 2, light,
chlorophyll, water, temp.)
* CO2 becoming limiting in clear sky, but light
limiting in cloudy days.
* Atmospheric CO2 is not limiting factor for C4
plants & submerged hydrophytes.
Introduction :
· Growth is a characteristic feature of all living
organisms.
· Growth is an irreversible increase in size of an organ or
its part or even of an individual cell
· Growth is accompanied by metabolic processes, that
occurs at the expanse of energy
PLANT GROWTH GENERALLY IS INDETERMINATE :
· In plants growth continues throughout their life. So it
is called as indeterminate or unlimited growth.
· Meristem continuously divides and add new cells to
the plant body, such activity of meristem is called as open
from of growth or indeterminate growth.
· Growth is diffused in animals but growth in plants is
localised. (Presence of meristem at certain locations in
plant body)
· Seed germination is the first step of plant growth.
Almost all the plants face a period of suspended growth
* If the suspension of growth is due to exogenously
controlled factors (environmental factors) then it is
called quiescence
· When the suspension of growth is due to the
endogenously controlled factors (hormonal, ge netic) then
it is termed as dormancy
GROWTH IS MEASURABLE :
At cellular level growth can be measured by measuring the
increases in the amount to protoplasm but it is very
difficult to measure directly, so growth is measured by a
variety of parameters, they are
(a) Increase in fresh weight
(b) Increases in dry weight
(c) Increase in surface area/ volume
(d) Increase in number of size of cells
Growth is measured by Auxanometer
PHASES OF GROWTH :
1. Cell division or cell formation or meristematic phase :
Number of cells is increased by cell division in this
phase. The cells of this region have rich protoplasm and
conspicuous nuclei, thin and primary cell wall &
abundant plasmodesmatal connections.
2. Cell enlargement or cell elongation phase : Size of cells
is increased due to vacuolation & TP (Turgor pressure)
and new cell wall depositions in this phase.
3. Cell maturation or differentiation phase : Cell wall
thickening and Protoplasmic modifications. Qualitative
changes in cells is important feature of this phase. It leads
to formation of mature tissues.
GROWTH RATE :
· Increased growth per unit time is termed as growth
rate.
· The growth rate shows an increase that may be
arithmatic or geometric
· Airthmatic-growth : In arithmatic growth only one
daughter cell among the two further divides while other
differentiates and become mature (stop dividing).
Ex. Root & shoot elongation of constant Rate. It is
methamatically expressed as
Lt = L0 + rt
where Lt : length at time ‘t’
L0 – length at time ;zero’
r – growth rate / elongation per unit time. It’s
curve is linear.
(b) Geometric Growth : Here both the progeny cells
following mitotic divisions retain the ability
to divide and continue to do so.
Ex. Early embryonic development/division in
zygote, division in unicellular organism. It is
mathematically represented as
W1 = W0ert
Where W1 = final size
(Weight, height, number etc.)
W0 = initial size at the beginning of period.
R = growth rate
In most system the initial growth is slow (lag phase) and it
increase rapidly thereafter at an exponential rate (log or
exponential phase), it is also called as “grand phase of
growth”. However with limited nutrient supply, the
growth slows down leading to a stationary phase or steady
phase. If we plot the parameter of growth against time,
we get typical for all cells, tissues and
organs of a plant.
r = relative growth rate and is also the measure of the
ability of the plant to produce new plant material referred
to as efficiency index.
Absolute and Relative growth rates :
Absolute growth rate :
Total growth which occurs in unit time in plant or plant
parts.
Relative growth Rate :
Total growth which occurs in unit time with respect to
initial parameter in plant or plant parts.
Relative growth rate is generally high in young developing
plant parts.
Time period - 7 days
* Both posses name absolute growth rate i.e 5 cm2 in 7
days.
* But high relative growth rate is in leaf ‘A’ about
100% while in leaf ‘B’ it is about 10%.
Conditions for growth
Water, oxygen and nutrient are very essential for
growth. The plant cells grow in size by cell enlargement
which in turn requires water. Turgidity of cells helps in
extension growth. Thus, plant growth and further
development is intimately linked to the water status of the
plant. Water also provides the medium for enzymatic
activities needed for growth.
Oxygen helps in releasing metabolic energy essential for
growth activities. Nutrients (macro and micro essential
elements) are required by plants for the synthesis of
protoplasm and act as source of energy.
DIFFERENTIATION, DEDIFFERENTIATION
AND REDIFFERENTATIONCells
Cell produced after division (mitosis) in meristem
transform of differentiate theirselves according to specific
function. This act leading to maturation of cells is called as
differentiation e.g. formation of primary permanent tissues
like parenchyma etc
The living and differentiated cells sometimes again have
to divide according to the requirements of plants for this
purpose these cells regain their dividing capacity, this
phenomenon is called as dedifferentiation e.g formation of
interfascicular cambium from the differentiated pemanent
parenchymatous cells and formation of cork cambium from
hypodermis.
 Such dedifferentiated cells divide and produce the cells
that once again lose the capacity to divide but mature to
perform specific functions, this is called as
redifferentiation. Example of redifferentiated tissues are
secondary xylem , secondary phloem, secondary cortex
etc.
Differentiation in plants is open , because cells/ tissues
arising out of the same meristem have different structures
at maturity and some of them have the ability to return to
division stage.
 The final structure at maturity of cell/tissue is also
determined by the location of the cell within for example .
Cells positioned away from root apical meristems
differentiate as root-cap cells, while those pushed to the
periphery mature as epidermis.
Plants follow different pathways in response to
environment or phases of life to form different kinds of
structures. This ability is called plasticity, e.g. heterophylly
in cotton, conriander and larkspur. In such plants, the
leaves of the juvenile plant are different in shape from those
in mature plants (internal plasticity) On the other hand,
difference in shapes of leaves produced in air and those
produced in water in buttercup also represent the
heterophyllous development due to environment
(Environmental plasticity). This phenomenon of
heterophylly is an example of plasticity
Development it considered as the sum of growth and
differentiation, which is controlled by both extrinsic
(environmental) and intrinsic (internal) factors
(i) Extrinsic factors – Light, temperature , water oxygen ,
nutrition
(ii) Intrinsic factors – Genetic factors (intracellular) and
PGR (intercellular)
PGRs are also called plant growth substance or plant
hormones or phytohormones.
PGRs are of two types-
(1) Plant growth promotors:
(i) Auxins (ii) Gibberellin (iii) Cytokinin

(2) Plant growth inhibitors :

(iv) Abscisic acid (v) Ethylene
Characteristic ''or'' chemical nature of plant growth
regulators :
• The plant growth regulators (PGRs) are small, simple
molecules of diverse chemical composition. They could be
(i) Indole compounds (indole-3-acetic acid = IAA) ± Auxin
(ii) Adenine derivatives (N6-furfurylamino purine =
kinetin) Þ Cytokinin
(iii) Derivatives of carotenoids (abscisic acid = ABA)
(iv) Terpenes (gibberellic acid = GA3) Gibberellin
(v) Gases (ethylene = C2H4).
AUXINS DISCOVERY -
• Term Auxin derived from Greek word 'auxein' which
means to grow.
• First of all Charles Darwin & Francis Darwin (son of
C. Darwin) observed the process of
• phototropism. They observed coleoptile bending in
Canary grass (Phalaris), responded to unilateral
illumination.
• After a series of experiments, it was concluded that the tip
of coleoptile was the site of transmittable influence (Auxin)
that caused the bending of the entire coleoptile. Auxin was
isolated by F.W. Went from tips of coleoptiles of oat seedling.
(First of all, Auxin was isolated from human urine)
• F.W. Went gave the name “auxin” to growth substance,
thus credit of discovery of auxin is given to F. W Went.
• Now IBA (Indone Butyric acid) have also been isolated
from plants (natural auxin) but IAA (Indole Acetic Acid) is
most widely occuring auxin in plants.
PHYSIOLOGICAL EFFECTS OF AUXIN –
(1) Apical Dominance (Characteristic function of
auxin) :- The phenomenon in which apical bud
dominates the growth of lateral buds is called Apical
Dominance.
Removed of shoot tips (decapitation) usually
results in the growth of lateral buds. It is widely
applied in tea plantations, hedge – making.
(2) Cell Division & Cell Enlargement : Mainly cell
elongation. Helps in cell division. Auxin also controls
the xylem differentiation.
(3) Root initiation :- Rooting on stem cuttings (widely
used in plant propagation) is promoted by IBA & NAA.
(4) Prevention of Abscission :- IAA, NAA prevent
premature abscission of plant organs. Promote the
abscission of older mature leaves & Fruits.
(5) Flower initiation :- Auxin promotes flowering in
Pineapple & Litchi plants.
(6) Parthenocarpy :- Seed less fruits can be produced by
spray of IAA. Eg. Tomato.
(7) Selective weed killer :- 2,4-D is widely used to kill
Dicot weeds, does not affect monocot plants. 2, 4-D + 2,
4, 5-T – Agent orange. It is used to prepare weed free
lawns.
Synthetic Auxins –
NAA – Napthalene acetic acid
2,4-D – 2, 4-Dichlorophenoxy acetic acid.
2,4,5-T – 2, 4, 5-Trichloro phenoxy acetic acid.
GIBBERELLINS
DISCOVERY –
First of all Japanese farmers observed symptoms of
the disease in rice seedings & called it bakanae disease
(Foolish seeding disease)
• Kurosawa and Sawada confirmed that rice plants
become thin, tall & pale due to infection of fungus
Gibberlla fujikuroi (Ascomycetes)
• Yabuta and Sumiki were first to extract a crystalline
substance from the Gibberella fungus to which they
named Gibberellin
• More than 100 type of Gibberellins (GA1,GA2GA3
….. GA100) are known. GA3 |C19H26O6] is
representative of all Gibberellins.
They are synthesized in buds, shoot, root and
germinating seeds.
PHYSIOLOGICAL EFFECTS OF GIBBERELLINS
–
1. Stem/ internode elongation (characteristic
function of gibberellins) :- Gibberellins induce
internode elongation just prior to flowering in Rosette
plants (Beet and Cabbage), this phenomenon is known
as Bolting effect .
2. Elongation of genetically dwarf plants :- When
gibberellins are applied to dwarf Maize, Pisum
etc.Plants then they become tall.

3. Seed germination :- Gibberellin induces the

synthesis of hydrolysing enzymes like amylase -
Lipases & Proteases in seed and these enzymes help in
seed germination by promoting breakdown of
endosperm.
4. Fruit & Flower enlarger: - Size of grape fruits
and length of stalk of stalk of bunch is increased by
GA Pomalin Þ GA + GK – used as apple enlarger.
5. In fermentation :- GA increase malt formation .
(Malting process) so helpful in brewing (wine)
industry.
6. Increase height of Sugarcane plant :- GA can
increase the yield in sugarcane upto 20 tonnes per acre.
7. GA delays the senescence in plant parts so the fruits
can be left on the tree longer so as to extend the market
period by application of GA.
8. Spraying juvenile conifers with Gas. Hastens the
maturity period, thus leading to early seed production
CYTOKININ (ck)
DISCOVERY-
• F.Skoog and his coworkers observed in tobacco
that callus can proliferate only if, in addition to auxin
the nutrient medium was supplemented with one of the
substances like extract of vascular tissues yeast extract,
coconut milk or DNA.
• Skoog and Miller later identified and crystellised
an active substance from autoclaved DNA of Herring
fish sperm, which stimulated cell division (mainly
Cytokinesis) They named this substance as kinetin.
Kinetin does not occur naturally in plants.
• The first natural cytokinin was identified &
crystalized from immature corn grains (corn krenes) and
coconut milk by Letham & named it as Zeatin.
• Cytokinins are derivative of Adenine nitrogen base
• Root tips are major sites of biosynthesis of CK. Also
synthesised in developing shoot buds, young fruits
etc.
Physiological effects and applications
(1) Cell division & Cell enlargement :- One of the most
important biological effect of CK (cytokinin) on plants
is induction of cell division. In tissue culture also.
(2) Formation of interfascicular cambium and induce
secondary growth.
(3) Morphogenesis :- Morphogenetic changes induced
by CK in presence of IAA.
High auxin + low CK - Root formation
(4) Counteraction of apical dominance :- promotes
growth of lateral buds.
(5) Breaking the dormancy of seeds :- Like GA the
dormancy of certain seeds can be broken by CK.
(6) Seed germination :- Seeds of parasite plant (Striga)
can germinate in the absence of host by CK treatment.
(7) Delay in senescence :- (Richmond Lang Effect) The
ageing process of leaves usually accompanies with
loss of chlorophyll & rapid catabolism. This is called as
senescence. senescence postponed by CK. (increase
short life of plant parts)

Parthenocarpy in some fruits.

(10) Pro-plastids modification.
(11) Phloem conduction (nutrients mobilisation)
(12) Femaleness.
(13) Flowering in SDP (also in long days).
(14) Induce stomatal opening :-
Bio–assay :–
(1) Tobacco pith cell division test
(2) Chlorophyll preservation (retension) test (delay in
senescence test)
(3) Soyabean and radish cotyledon cell division test.
ABSCISIC ACID (ABA)
DISCOVERY-
First Growth inhibitor was identified & isolated by Bennet
Clark (1953) and Kefford from dormant Potato tuber and
called Inhibitor –B.
• Addicott & Okhuma isolated a substance from mature
cotton fruits and named it as Abscission-II
• Waring & Robinson isolated a growth inhibitor from
old Betula leaves & called it Dormin.
• Later it was proved that inhibitor- B , Abscission -II &
Dormin are same and they were regarded as Abscisic
acid.
• ABA is mainly synthesised in senescent organs, old
leaves
• ABA is also known as stress hormone because it
protects plants from adverse conditions like water stress by
inducing closing of stomata. ABA increase tolerance of
plants to various type of stresses.
• Common growth inhibitor in plants is ABA
PHYSIOLOGICAL EFFECT OF ABA -
• Induces senesence and abscission - ABA causes
senescence and abscission of leaves and fruits by
increasing the activity of cellulase & pectinase enzymes.
• Induces Bud & Seed dormancy - ABA increases bud &
seed dormancy (inhibits seed germination).
• Stomatal closing -ABA closes stomata under the water
stress condition (Anti transpirat).
• In most situations ABA act as an antagonist of GAs
(Anti GA)
ETHYLENE
DISCOVERY
Ethylene can be included in both growth promotor
and growth inhibitor but mainly it is a growth inhibitor
hormone
• Ethylene is simple gaseous PGR/ It is a hydrocarbon
reported as a fruit ripening hormone.
• H.H. Cousins confirmed the release of volatile
substance from ripened oranges that hastened the ripening
of stored unripened bananas. Later this volatile substance
was indentified as ethylen a gaseous PGR.
• Biosynthesis of Ethylene takes place by methionine
amino acid/ Ethylene is synthesised in large quantity by
ripening fruits and senescent organs.
PHYSIOLOGICAL EFFECTS OF ETHYLINE -
1. Post harvest ripening of fruits :- Citrus, Oranges ,
Banana, Apple, Tomato etc. Now a days Ethephon/
CEPA (Chloroethyl Phosphonic acid) is used at
commercial level as source of ethylene. Ethylene
enhances the respiration rate during ripening of the
fruits. This rise in respiration is called respiratory
climactic.
2. Stimulation of senescence & abscission of flower and
fruit. Thinging of cotton, cheryy, walnut.
3. Flowering and synchronising fruting in pineapple.
Flowering in mango.
4. Triple response on stem :-
(i) Apical hook formation in dicot seedings
(ii) Swelling of axis
(iii) Horizontal growth of seedings
5. Promotes root growth :- Ethylene promotes root
growth and also stimulates the formation of root hairs,
thus helping the plant to increase their absorption
surface.
6. Femaleness (Feminising effect) in cucumbers.
(Promotes female flowers to increase the production)
7. Ethylene breaks seed and bud dormancy. Initiates
germination in peanut seeds, sprouting of potato tubers.
8. Ethylene promotes rapid internode/ petiolde
elongation in deep water rice plants . It helps leaves/
upper parts of the shoot to remain above water.
PHOTOPERIODISM
• Effect or requirement of relative length of day
(photoperiod) & night (dark phase) on flowering of
plants it called as photoperiodism
• The phenomenon of photoperiodism was first
discovered by Garner & Allard on Maryland mammoth
(a mutant variety of Tobacco) and Biloxy soyabean.
• Garner & Allard classified the plants in following
groups
1. SDPs (Short Day Plants):-
* These plants flower on exposure to photoperiod
shorter than their Critical day length.
* In SDPs the dark period is critical and must be
continuous. Thus SDP are also called as LNP (long Night
plants) if this dark period is interrupted even with a brief
exposure of light , the SDP will not flower.
* Examples of SDPs :-Tobacco, Soyabean, Viola,
Xanthium (Cocklebur), Chrysanthemum, Cannabis,
Coleus, Chenopodium, Musturd, Dahila, Sugarcane,
Strawberrty, Cosmos. Rice, Bryophyllum etc.
2. LDPs (long Day Plants) :-
* These plants flowers when they are exposed to
photoperiod longer than their critical dya night.
* A brief exposure of light in the dark period stimulates
flowering in LDPs.
* The light period is critical for LDPs.
* Gibberellin can induce flowering in LDP even under
non-inductive photo period condition.
Ex. Henbane (Hyoscyamus), Spinach, Sugarbeets
,Radish, Carrot, Wheat, Larkspur , Barley, Avena,
Potato.
3. DNPs (Day Neutral Plants) :-
* These plants do not need a specific light period for
the flowering
Ex. Maize, Cotton, Tomato, Sunflower, Cucumber
Phytochrome :
* A light sensitive pigment phytochrome is responsible
for flowering induction and present in leaves.
* Phytochrome exists in two different forms
* Pr (Phytochrome Red) Pfr (Phytochrome Far Red)Pfr
(Phytochrome Far Red)
* & Both forms of phytochromes are photobiochemically
interchangeable into each other
 Phytochrome Pfr (P230) is active form which controls
many photophysiological processes in plants like -
Flowering, seed germination ect.
* Stimulation of critical photoperiod is perceived by
leaves
* Florigen hypothetical hormone (not isotaled yet from
plants) which migrates from leaves to shoot apices for
inducing flowering
VERNALISATION OR YAROVIZATION
* Effect of low temperature on the initiation and
development of flowers, was first realised by Klippart
(Experiments on winter wheat and Spring wheat)
* Mainly embryo tip, shoot apex & leaves perceive
stimulus of low temperature.
* Concept of hormone Vernalin in vernalisation was
given by Melcher. This is a hypothetical plant hormone
because has not been isolated
* Vernalisawtion of seeds or plant propagules in
laboratory can be induced at 1°C to 10°C in presence of
O2 & H2O.
* If vernalized plant propagules are kept at high
temperature then effect of vernalisation is reversed, this
is called devernalization.
* Two types of varieties are found in wheat, barley and
rye (cereal) winter and spring variety. Winter varieties
require vernalisation whereas spring varieties do not.
* Sugar beet, cabbage, carrot etc. are biennial plants.
* Significance :-
(i) Better & early flowering
(ii) Prevention of precocious reproductive
development late in the growing season.
(iii) Shortening the life span of the plants.
PLANT SENESCENCE
* Period from maturity to death of an organ or plant is
known as senescence.
* During the senescence, higher rate of catabolism is
started, under the control of growth hormones like ABA
and ethylene . Senescence occurs as a result of ageing and
leads to the death of plant parts or whole plant. (Study of
senescence and ageing is phyto- Gerontology)
ABSCISSION
* Detachment of senescent or mature plant organs like
leaves, fruits, Flowers due to change in hormonal activity.
* There is the formation of separation layer (Abscission
layer) at the region of attachment of these parts. Cell wall
layers and middle lamella are dissolved by the activity of
cellulase and pectinases enzymes during the abscission
* Hormone ABA is main controller of abscission process.
* Respiration is a Amphibolic & exergonic cellular
process.
* Respiration is an enzymatic process, which is also
known as internal respiration / tissue respiration/dark
respiration / cellular respiration / mitochondrial
respiration.
* An important feature of respiration is liberation of
metabolic energy as ATP.
Respiratory substrate :
(Carbohydrates - Fats - Protein - others)
* When respiratory substrates are carbohydrates like
glycogen, starch, sucrose, hexose or fats, then
respiration is known as floating respiration.
* When protein is oxidised in respiration, then
respiration is Known as protoplasmic respiration
protoplasmic components or cellular proteins may
oxidised at the time of starvation & disease.
* Exceptionally oxidation of proteins in legume seeds
is called floating respiration.
Types of respiration :
(A) Aerobic respiration :- The complete oxidation of
food with the use of oxygen and when entire
carbon released, as CO2 is called as aerobic
respiration.
(B) Anaerobic respiration :- This is an incomplete
oxidation.
* When food is oxidized into alcohol or organic acids
without use of oxygen.
During it most of the energy is lost in form of heat. It
occurs in cytoplasm and only 2ATP are produced.
* Anaerobic respiration may takes place in bacteria,
some lower parasitic animals (Ascaris, Taenia) plants,
R.BCs. & muscles of human body. When oxygen is not
available, then food is incompletely oxidised in to some
organic compounds like ethanol, acetic acid, lactic acid.
* In muscle cells & some bacteria, the energy is
produced by breaking of glucose into lactic acid inside
the cells.
* The amount of energy released in anaerobic
respiration is much less than aerobic respiration.
* Fermentation is performed by only some fungi &
some bacteria (only by microbes) and is an
extracellular process. There is gain of 2ATP.

* Both anaerobic respiration and fermentation are

incomplete oxidations.
* Inhibitory effect on respiration high conc. of
oxygen is called Pasteur effect.
Types of Respiration
(1) Glycolysis – Occurs in cytosol/cytoplasm
(2) Formation of Acetyl COA – (Link Reaction)
Perimitochondrial space (outer chamber)
(3) TCA cycle or Kreb's cycle – Matrix of mitochondria
& cytosol in bacteria
(4) ETS – Occurs in cristae or inner membrane of
mitochondria and Oxidative phosphorylation –
Occurs in Oxysome head (F1 particle)
(1) Glycolysis – EMP – (Embden, Meyerhof,
parnas) pathway.
* The glycolysis is common phase for aerobic &
anaerobic respirations both.
* Glycolysis involves a series of ten biochemical
reactions in cytoplasm.
* In glycolysis, neither consumption of oxygen nor
liberation of CO2 take place.
* In glycolysis, 1 glucose, produces 2 mol. of
pyruvic acids (3C)
* 2NADH2 & 2ATP are generated in glycolysis, which
are equal to 8 ATP.
* Substrate level phosphorylation forms 4 ATP :-
[When the substrate releases energy for
phosphorylation of ADP OR formation of ATP, without ETS
then called as substrate level phosphorylation]
* Glycolysis is also known as oxidative anabolism or
catabolic resynthesis, because it links with anabolism
of fats and amino acids. an intermediate PGAL is used
for the synthesis of glycerol later forms fats or lipid. PGA
is used for synthesis of Serine, Glycine, Cystine. Alanine
forms from pyruvate.
* 1, 3, 10 are irrev. reactis in EMP pathway.
(2) Formation of Acetyl-Co-A :- (Link/Gateway reaction)
* When respiration is aerobic, then pyruvic acid is
oxidised to form 2C compound – Acetyl Co-A. It occurs
in presence of O2 and CO2 is released first time during it.
* Acetyl Co-A is a connecting link between glycolysis &
Krebs-cycle. Decarboxylation and dehydrogenation
(Oxidative decarboxylation) take place during formation of
acetyl Co-A.
* Acetyl Co-A is formed in mitochondrial matrix by
enzyme pyruvate dehydrogenase complex. (Mg+
+
, LA (Lipoic Acid), TPP(Thiamine
pyrophosphate), NAD, CoA)
* Acetyl Co-A is also common intermediate between
fat & carbohydrate metabolism.
(3) Kreb cycle / TCA (Tricarboxylic acid) Cycle /
Citric acid cycle :-
* This cycle was discovered by H.A. Kreb. (Nobel
prize)
* TCA cycle occurs in mitochondrial matrix or power
house of cell.
* Kreb cycle begins by formation of citric acid
[TCA(Tri carboxylic acid)] & O.A.A. is the acceptor
molecule of Acetyl CoA in Kreb's cycle.
* A number of Krebs cycle intermediates are used
in synthetic (anabolic) pathways, thus TCA cycle
is also called amphibolic pathway or
anaplerotic pathway.
* Succinyl CO–A is important for synthesis of
porphyrin ring compounds like Chlorophylls,
Phytochromes, Phycobilins, Cytochromes,
Haemoglobin etc.
* a-ketoglutaric acid (5c) involves in Amino Acid
formation (Nitrogen-metabolism)
* Oxidation occurs at 4 sites in Kreb cycle.
* 3NADH2, 1FADH2 & 1GTP (ATP) produced by
each turn of TCA cycle.(=12 ATP)
* All the enzymes of TCA cycle, except marker
enzyme Succinic dehydrogenase (on inner
mitochondrial membrane) present in matrix.
ETS & Oxidative Phosphorylation (Terminal
Oxidation of NADH2 & FADH2)
All the reduced hydrogen acceptors like NADH +
H+ and FADH2 move the ETS where they release
their hydrogen and get reoxidised to NAD+ & FAD,
so that they can again enter into the respiration
process.
Cytochromes are cyt.b, cyt.-C1, cyt.-C, cya-a & cyt-a3.

Now components of ETS and ATP synthase are

categorised as follows
Name of Components of
Complexes ETS
Complex-I FMN, Fe-S NADH
dehydrogenase
Complex-II Succinate dehydrogenase,
FADH2
Complex-III Cytochrome b Cyt C1
Complex IV Cy. A and cyt. a3 , 2Cu centres
( Cytochrome oxidase)
Complex V ATP synthase / ATP pase
* CoQ/UQ is a mobile I (e– +H+) carrier and cyt- c is
mobile e carrier UQ is located within the inner
mitochondrial membrane, cyt-c is a small protein
attached to the outer surface (toward PMS) of the inner
membrane.
O2 is last e– acceptor in oxidative phosphorylation & due
to its reduction water is formed

During each step of mitochondrial ETS , redox reaction

occurs and energy is released which is utilized in creation
of proton gradient for the synthesis of ATP in presence of
oxygen (Oxidative phosphorylation)
* During the ETS, NADH + H+ gives it’s 2e–/2H+ to
FMN I the respiratory chain, thus 3 ATP are
generated, while FADH2 give it’s 2e-/2H+ to CoQ hence
only 2 ATP are formed during the process of oxidative
phosphorylation
Enzyme cytochrome oxidase is responsible for oxidation
of cyt a3 & reduction of O2. (Transfer of electrons from
cytochrome a3 to O2)
Cyanide (CN) inhibits the activity of cytochrome oxidase
(inhibits the oxidation of cyt-a3 & reduction of O2)
In mitochondria of some plants alternative oxidase
system is present in which ETS continues even in the
presence of cyanide. This type of respiration is known as
cyanide resistant respiration of Alternate electron
pathway . Ex. Spinach, Pea.
Chemiosmotic theory/ Coupling theory of Oxidative
phosphorylation:
During ETC of respiration FMN & CoQ can releases H +
ions in perimitochondrial space and leads to differental
H+ ion concentration across inner mitochondrial
membrane. This differential H+ ion concentration across
inner mitochondrial membrane leads to creation of
proton gradient (pH gradient) and Electrical potential
(difference in charge) Both are collectively known as
Proton motive force (PMF).
PMF does not allow stay of H+ ions in Perimitochondrial
space (PMS) so they return towards the matrix through
F0 particles selectively . (Facilitated diffusion)

Passage of 2H+ through F0 particle or coupling factor or

proton channel leads to synthesis of 1 ATO
(i) Glycerol phosphate shuttle :- In brain/muscle
cells.
(ii) Malate aspartate shuttle :
* In prokaryotes, shuttle mechanism is absent.
They always get 38 ATP from aerobic
respiration of 1 glucose mol.
* Cyanide inhibits the activity of cytochrome
oxidase & inhibits the oxidation of cyto-a3.
* In mitochondria, of some plants alternative
oxidase system is present, in which ETS
continues even in presence of cyanides. This type
of respiration is known as cyanide resistance
respiration or Alternate electron pathway. Ex.
Spinacea, Pisum.
ETC
Oxidative phosphorylation
Chemiosmotic theory / Coupling theory :
* During ETC of respiration CoQ & FMN can
releases H+ ions in perimitochondrial space and
leads to differenctial H+ ion concetration across
inner mitochondrial membrane. This differential H+
ion concentration across inner mitochondrial
membrane leads to creation of proton gradiant
(PH gradient) and Electrical potential (diffrence of
charge). Both are collectively known as Proton
motive force (PMF).
* PMF donot allow stay of H+ ions in
Perimitochondrial space (PMS) so they return
towards the matrix through F0 particles
selectively.
* The passage of 3H+ ions activate ATP synthase
and gives rise to 1ATP from ADP & Pi.
* Some physiologist believe that passage of 2H +
ions through F0 particle or coupling factor or
proton channel leads to synthesis of 1 ATP.
(B) Anaerobic respiration :- This is an incomplete
oxidation.
* When food is oxidized into alcohol or organic
acids without use of oxygen.
During it most of the energy is lost in form of
heat. It occurs in cytoplasm and only 2ATP are
produced.
Lactic acid anaerobic respiration
Alcoholic fermentation: In yeast
Value of R.Q. depends upon the type of respiratory
substrate used & measured by Ganong's
respirometer.
* Fatty seeds germination-castor, R.Q. < 1
* At the time of formation of fatty seeds, (maturing
fatty seed) R.Q. is more than 1 because more
CO2 is released than O2 consumed.
 Minerals & Nutrition
WATER RELATIONS -Theory -
Generally all living organism have same basic needs.
They require macromolecules, such as carbohydrates,
proteins and fats, and water and minerals for their
growth and development.
METHODS TO STUDY THE MINERAL,
REQUIREMENTS OF PLANTS
In 1860, Julius von Sachs, a prominent German
botanist, demonstrated for the first time, that plants
could be grown to maturity in a defined nutrient
solution in complete absence of soil.Since then, a
number of improvised methods have been employed
to try and determine the mineral nutrients essential
for plants. This technique of growing plants in a
nutrient solution is known as Hydroponics.
The essence of all these methods involves the
culture of plants in a soil-free, defined mineral
solution. These methods require purified water and
mineral nutrient salts.

After a series of experiments in which the roots of

the plants were immersed in nutrient solutions and
wherein an element was added/removed or given in
varied concentration, a mineral solution suitable for
the plant growth was obtained.
By this method. essential elements were identified and
their deficiency symptoms discovered.

Hydroponics has been successfully employed as a

technique for the commercial production of
vegetables such as tomato, seedless cucumber and
lettuce.

It must be emphasized that the nutrient solutions

must be adequately aerated to obtain the optimum
growth.
Diagrammatic views of the hydroponic technique is given
in Figure.
Essential Mineral Elements
Most of the minerals present in soil can enter
plants through roots. In fact, more than sixty
elements of the 105 discovered so far are found
in different plants.

Some plant species accumulate selenium, some

others gold, while some plants growing near
nuclear test sites take up radioactive strontium.
There are techniques that are able to detect the
minerals even at a very low concentration (10 8
g/ml). The question is, whether all the diverse
mineral elements present in a plant, for example,
gold and selenium as mentioned above, are really
necessary for plants? How do we decide what is
essential for plants and what is not ?
Figure :- Hydroponic plant production.
(i) Plants are grown in a tube or trough placed on slight incline. The
solution flows down the tube and returns to the reservoir due to
gravity.
(ii) A pump circulates a nutrient solution from a reservoir to the
elevated end of the tube. Inset shows a plant whose roots are
continuously bathed in aerated nutrient solution. The arrows indicates
the direction of the flow.
Criteria for essentiality
Mentioned below is the criteria for essentiality
of an element:
(a) The element must be absolutely necessary
for supporting normal growth and
reproduction. In the absence of the element
the plants do not complete their life cycle or
set the seeds.
(b) The element must be directly involved in the
metabolism of the plant.
(c) The requirement of the element must be
specific and not replaceable by another element.
In other words, deficiency of any one element
cannot be met by supplying some other element.

According to the above criteria only a few

elements have been found to be exactly essential
for plant growth and metabolism. These elements
are further divided into two broad categories
based on their quantitative requirements.
(i) Macronutrients, and
(ii) Micronutrients
1. Macronutrients are generally present in plant
tissues in large amounts (in excess of 10 m
mole Kg-1 of dry matter). The macronutrients include
carbon. hydrogen, oxygen, nitrogen,
phosphorous, sulphur, potassium calcium and
magnesium. Of these, carbon, hydrogen and
oxygen are mainly obtained from CO2 and H2O,
while the others are absorbed from the soil as
mineral nutrition.
2. Micronutrients or trace elements, are needed
in very small amounts (less than 10 m mole
Kg-1 of dry matter). These include iron,
manganese. copper, molybdenum, zinc, boron,
chlorine and nickel.

3. In addition to the 17 essential elements named

above, there are some beneficial elements
such as sodium, silicon, cobalt and selenium.
They are required by higher plants.
ESSENTIAL MINERAL ELEMENTS
Most of the minerals present in the soil can enter
plants through their roots. More than sixty mineral
elements out of 105 discovered so far have been
recorded in different plants. We can detect the presence
of minerals even at a very low concentration (10–8 g.mL)
in plants.
Some plants accumulate heavy and toxic minerals
such as gold and selenium from the soil. While some
plants growing near the nuclear test sites take up
radioactive strontium. But , all the diverse mineral
elements found in plants are not essential for plants.
An element is said to be essential for plants if it
has a specific structural or physiological role and
without which plants cannot complete their life
cycle. The criteria to know the essentiality of an
element was proposed by Arnon and Stout in
1939.
Criteria for Essentiality
The criteria to determine the essentiality of an
element include the following aspects
1. The element must be absolutely necessary for
supporting normal growth and reproduction of
plants. In the absence of the element, the plants
do not complete their life cycle or set the seeds.
2. The element must be directly involved in the
metabolism of the plant. It should form a
component of either a structural or functional
molecule.
3. The requirement of the element must be
specific and not replaceable by another element.

4. Absence or reduced availability of the element

causes disorders.
5. The disorders caused by absence or
deficiency of an element can be corrected only
by the availability of that element.
Based upon the above criteria, 17 elements have
been found to be essential for plant growth and
metabolism. They are C,H,O,N,P,K, Ca, Mg,
S,Fe,B, Mn,Cu,Zn, Mo,Cl and Ni. Others are called
non-essential elements.
Classification of Essential Elements
A. On the basis of quantitative requirement : The
essential elements (nutrients) are divided into two
broad categories based on their quantitative
requirements. These are
1. Macronutrients
2. Micronutrients
1. Macronutrients : These are present in large amounts
in plant tissues, i.e., in excess of 10 m mole kg–1 of dry
matter.
Macronutrients are easily detectable because they
are present in large quantity. They are usually involved in
the synthesis of organic molecules
These macronutrients are nine in number - Carbon,
hydrogen oxygen, nitrogen, phosphorus, sulphur
potassium, calcium and magnesium. Out of these, carbon,
hydrogen and oxygen are mainly obtained from CO2 and
H2O, while the others are absorbed from the soil as
mineral nutrition.
2. Micronutrients or trace elements: These are
the essential elements which are required by
plants in small amounts, less than 10 m mole kg–1
of dry matter Micronutrients are mostly involved
in the functioning of enzymes. Since,
micronutrients are required in traces, they are
also called trace elements.
Micronutrients are 8 in number.
Iron, manganese, copper, molybdenum, zinc,
boron, chlorine, nickle.
In addition to the 17 essential elements named
above, there are some beneficial elements such
as sodium, silicon, cobalt and selenium. They are
required in metabolic activities of higher plants.
B. On the basis of Function : Essential elements can
also be grouped into four broad categories on the
basis of their diverse functions. These categories are :
1. Structural elements : Some essential elements
are components of biomolecules and hence are the
structural elementsof cells. They are carbon,
hydrogen, oxygen and nitrogen. For example,
(i) C, H and O are components of cellulose and
many other biomolecules.
(ii) Nitrogen is a constituent of all amino acids,
proteins, chlorophyll etc
2. Components of energy-related compounds :
Some essential elements become components of
energy related chemical compounds in organisms
For example'
(i) Magnesium is a component of chlorophyll that
takes part in conversion of light energy into
chemical energy.
(ii) phosphorus is a component of ATP which
functions as energy currency of the living systems.
3. Enzyme activators or inhibitors : Many essential
elements are responsible for activating or inhibiting
enzymes. For example,
(i) Mg2+ is an activator for both ribulose bisphosphate
carboxylase – oxygenase (RuBisCO) and
phosphoenol pyruvate carboxylase (PEPCase),
both of which are critical enzymes involved in
carbon fixation during Photosynthesis
(ii) Zn2+ is an activator of alcohol dehydrogenase and
Mo is an activator of nitrogenase during the
nitrogen metabolism.
(iii) Mn2+ is involved in photolysis of water.
4. Maintaining osmotic potential : Most of the
osmotic potential of cell sap is due to inoganic salts
such as nitrate, Cl–,K+,sulphate etc. Osmostic potential
is required for water absorption and maintenance of
cell turgidity.
Apart from this, mineral elements perform other
important functions such as :
(i) Movements : Potassium plays an important role in
the opening and closing of stomata and other turgor
movements.
(ii) Buffer activity : Phosphates, weak acids and their
salts possess buffer activity against changes in pH.
Role of Macro and Micronutrients
As we have discussed, essential elements
perform several functions. They participate in
various metabolic pro cesses in the plant cells such
as permeability of cell membrane, maintenance of
osmotic concentration of cell sap, buffering action,
enzymatic activity and act as major constituent of
macromolecules and coenzymes. Let us now study
the various forms and functions of essential mineral
elements in detail.
1. Nitrogen : This is the mineral element required
by plants in the greatest amount.
Absorption : It is absorbed mainly as NO3– from
the soil. it can also be taken up as NO2– or NH4+.
Requirement : Nitrogen is required by all parts
of a plant, particularly the meristematic tissues
and the metabolically active cells.
Physiological role : Nitrogen is one of the
major constituent of proteins, nucleic acids,
vitamins and hormones.
Being a component of such a large number of
biomolecules, nitrogen is essential for all types of
metabolic activities, photosynthesis, respiration, cell
growth, cell division and reproductive growth.
2. Phosphorus :
Absorption : It is absorbed in the form of
phosphate ions (either as H2PO4– or HPO42–) from
the soil.
Requirement : Inside the plant, phosphorus is
stored in developing fruits, seeds, storage organs
and young meristematic tissues.
Physiological role :
(i) Phosphorus is a constituent of cell membranes,
certain proteins, nucleotides, nucleic acids
(DNA and RNA), ATP, NADP and nucleoproteins.
(ii) It is required for all phosphorylation reactions.
ATP contains energy-rich phosphate bonds.
(iii) Its deficiency causes purple or red spots on
leaves and delay in seed germination
3. Potassium :
Absorption : It is absorbed as K+.
Requirement :In plants, it is required in more
abundant quantities in the meristematic tissues,
buds, leaves and roottios.
(i) lt helps to maintain an anion-cation balance in
cells.
(ii) lt is involved in protein synthesis.
(iii) lt helps in opening and closing of stomata.
(iv) lt is responsible for activating several enzymes
connected with phosphorylation, photosynthesis,
respiration, synthesis of chlorophyll etc.
(v) It helps in maintaining turgidity of the cells.
(vi) lts deficiency causes scorched leaf tips, loss of
cambial activity and plastid disintegration.
4. Calcium :
Absorption : It is absorbed in the form of
calcium ion (Ca2+) from the soil.
Requirement : Calcium is required by
meristematic and differentiating tissues. It
accumulates in older leaves.
Physiological role :
(i) lt is a required for synthesis of cell wall
during cell division, particularly as calcium pectate
in the middle lamella.
(ii) It is also required during the formation of mitotic
spindle and organisation of chromosornes.
(iii) It is involved in the normal functioning of the
cell membranes.
(iv) It activates several enzymes like ATPase,
phospholipases, a-amylase etc.
(v) It plays an important role in regulating
metabolic activities.
5. Magnesium :
Absorption : It is absorbed by plants in the form
of divalent magnesium ion (Mg2+).
Requirement : It is required in growing areas of
root and stem, seeds, leaves etc.
Physiological role :
(i) It occurs as magnesium pectate in middle lamella.
(ii) It is a constituent of the ring structure of
chlorophyll
(iii) It is required for binding of ribosome subunits
during protein synthesis.
(iv) It activates several enzymes involved in
photosynthesis and respiration.
(v) It is involved in the synthesis of DNA and RNA.
6. Sulphur :
Absorption : Plants absorb sulphur from the
soil in the form of sulphate ions (SO42–)
Requirement : It is required by young leaves
and meristems.
Physiological role :
(i) It is a constituent of two amino acids methionine
and cysteine.
(ii) It is also used in the synthesis of some vitamins
(thiamine and biotin), coenzyme-A and ferredoxin;
which are involved in various metabolic activities.
(iii) Its deficiency causes accumulation of
anthocyanin.
7. Iron : It is required in larger amounts in
comparison to other micronutrients
Absorption : Plants obtain iron in the form of
ferric ions (Fe3+)
Requirement : Everywhere in the body of plants
Physiological role :
(i) It is an important constituent of proteins involved in
the transfer of electrons like ferredoxin and
cytochromes. It is reversibly oxidised from Fe 2+ to Fe3+
during electron transfer in photosynthesis and
respiration.
(ii) It activates catalase and some other enzymes.
(iii) It is essential for the formation of chlorophyll and
other pigments
8. Manganese :
Absorption : It is absorbed in the form of
manganous ions (Mn2+) from the soil.
Requirement : Leaves and seeds
Physiological role :
(i) It activates several enzymes involved in
photosynthesis, respiration and nitrogen
metabolism.
(ii) It plays a major role in the splitting of water to
liberate oxygen, during photosynthesis.
(iii)Its deficiency causes grey spots in oats.
9. Zinc :
Absorption : Plants absorb zinc from soil as Zn2+
ions.
Requirement : Everywhere in the body of plants.
Physiological role :
(i) It activates various enzymes, especially
carboxylases.
(ii) It is also required in the synthesis of auxin.
(iii) It has role in tryptophan synthesis and its
deficiency produces leaf malformation or little leaf.
10. Copper :
Absorption : Plants absorb copper as cupric ions
(Cu2+) from the soil.
Requirement : Everywhere in the body of plants.
Physiological role :
(i) It is essential for the overall metabolism in plants.
(ii) Like iron, it is associated with certain enzymes
involved in redox reactions.
(iii) It is reversibly oxidised fron Cu+ to Cu2+ [ Cu+
Cu2+ + e–] during electron transport. Hence, copper is
essential for both photosynthesis and respiration.
11.Boron :
Absorption : It is absorbed as BO33– or B4O72–
Requirement : It lt is required in leaves and
seeds.
Physiological role :
(i) It is required for uptake and utilisation of Ca2+.
(ii) It is involved in functioning of cell membrane
and pollen germination.
(iii) It helps in translocation (transport) of
carbohydrates through phloem.
(iv) It is responsible for cell elongation and cell
differentiation.
(v) Its deficiency causes loss of apical dominance
and absence of root nodules in legumes.
12. Molybdenum :
Absorption : Plants absorb it from the soil in the
form of molybdate ions (MoO22+)
Requirement : Everywhere in the body of plants,
however, it is more commonly found in roots
Physiological role : It is a component of several
enzymes, including nitrogenase and nitrate reductase,
both of which participate in nitrogen metabolism. lt is
essential for nitrogen-fixation.
13. Chlorine :
Absorption : Chlorine is absorbed in the form of
chloride anion (Cl-) from the soil.
Requirement : Everywhere in the body of plants.
Physiological role :
(i) Alongwith manganese, chlorine is involved in
liberation of oxygen during photolysis of watetr in
photosynthesis.
(ii) Alongwith Na+ and K+, It helps in determining the
solute concentration and the anion-cation balance in
cells.
14. Nickel :
Absorption : It is absorbed from the soil as a
divalent cation (Ni2+)
Requirement : Leaves and roots
Physiological role :
(i) It is a component of two enzymes, urease and
hydrogenase
(ii) It is involved in the metabolism of urea.
MECHANISM OF ABSORPTION OF ELEMENTS
Soil is the main sources of mineral salts. These
mineral salts are mainly absorbed by the (Sub
terminal) meristematic region of the roots. Also by
root hair zone. Phosphate mainly absorbed by root
hairs.

The movement of mineral ions is usually called as

flux. The inward movement inside the cell is called
influx and outward movement is effux.
The mineral absorption occurs mainly in two phases –

In first phase, an initial rapid uptake of ions into the or

Outer space of the cell occurs, the apoplast
(intercellular spaces and cell wall), which is a passive
process.
In the second phase of uptake, the ions are taken
in slowly into the inner space, the symplast of the
cell which is both active and passive process
(mainly active process). The passive movement of
ions in symplast occurs through ion channel which
are trans-membrane proteins and act as selective
pores. The active influx and efflux from symplast
occurs with the help of pump proteins and with
expenditure of energy/ ATP.
Methods of Mineral Absorption –
(A)Passive (B)Active
(C) Passive absorption of Minerals : (Without
expenditure of ATP)
1. By simple diffusion : According to this method
mineral ions may diffuse into root cells from the soil
solution
2. By mass flow : According to this method mineral ions
absorption occurs with flow of water under the
influence of transpiration pull.
Why absorption is not completely passive ?
1. Minerals are present in the soil as charged
particles (ions) which mostly can not move across
cell membranes passively.
2. The concentration of minerals in the soil is usually
lower then the concentration of minerals in the
roots.
Active ion absorption : (By expenditure of ATPs)
In this process ions enter into more
concentrated cell sap from less concentrated
soil solution by consumption of energy from
ATP.

Specific proteins in the membrane of root hairs

actively pump ions from the soil into the
cytoplasm of the epidermal cells.
Like all cells, endodermal cells have many
transport proteins embedded in their plasma
membrane ; they let some solutes cross the
membrane, but not others. Transport proteins of
endodermal cells of root are control points , where
a plant adjust the quantity and types of solutes that
reach the xylem. The root endodermis because of
the layer of suberin has the ability of actively
transportations in one direction only.
TRANSLOCATION OF MINERAL IONS/ SOLUTES
When ions reach to the xylem of root by active or
passive absorption or by cumulative activity of both
then transport of these ions towards stem and all
the parts of plant occurs due to transpiration flow
through xylem
Main storage regions or sinks for minerals
elements are growing regions of the plants like
apices and lateral meristem, young leaves,
developing flowers, fruits, seeds and storage
organs.
Analysis of xylem sap shows that some nitrogen is
translocated in the form of inorganic ions and most
of it is translocated in the form of organic amides.

Like this small amount of phosphorus and sulphur

are also translocated in the form of organic
compounds. Except this small amount of materials
are also transferred between xylem and phloem.
So we cannot distinctly differentiate that xylem only
translocate inorganic nutrients and phloem
translocate only organic materials , as was
believed before.
METABOLISM OF NITROGEN INTRODUCTION
1. Nitrogen is an essential element in all living
organism. It is the constituent of amino acids,
proteins, hormones chlorophylls and many of the
vitamins.
2. Nitrogen is present in the atmosphere
abundantly in N2 form, but eukaryotes and many
prokaryotes can’t uptake nitrogen directly from
the atmosphere. The nitrogen enter into the soil
from atmosphere by fixation process. Then from
the soil plants absorb it and from plants it moves to
animals and fulfil the requirement of all living
beings.
3. Plants competes with microbes for the limited
nitrogen that is available in the soil, so nitrogen is
a limiting nutrient for both natural and agricultural
ecosystem
The nitrogen cycle showing relationship between the
three main nitrogen pool- atmosphere, soil and
biomass.
Steps of N2 Cycle :
1. N2 fixation
2. Uptake and assimilation of nitrogen by plants
3. Ammonification
4. Nitrification
5. Denitrification
Nitrogen Fixation
Process of conversion of atmospheric N 2 into
nitrogenous compounds (NH3,NO3–) is called
nitrogen fixation
Nitrogen fixation is of two types :
1. Abiological / Physicochemical
2. Biological
Abiological Nitrogen Fixation
A biological Nitrogen Fixation / Physicochemical N 2
fixation is further divided into two types
(A) Atmospheric or electrical
(B) Industrial
(A) Atmospheric or electrical : In nature
lightening and ultraviolet radiation provide enough
energy to convert nitrogen to nitrogen oxides
(NO, NO2, N2O).
(B) Industrial N2 fixation : In the presence of
high pressure , temperature and catalysts at
morphemic nitrogen and hydrogen combines to
form ammonia (NH3). This ammonia is used in the
formation of chemical fertilizers.
Biological Nitrogen Fixation/ Diazotrophy
Conversion of atmospheric nitrogen (N 2) into
inorganic nitrogenous compounds like – NH 3 by
living organisms is called biological nitrogen fixation
or Diazotrophy.

NITORGEN Fixing organisms (Diazotrophs)-

(A) Free living or- symbiotic
Eubacteria :- Azotobacter, Beijernickia, (both
aerobic) and Clostridium, Rhodospirllum (both
anaerobic)

Cyanobacteria (Blue green algae) – Nostoc,

Anabaena, Tolypothrix, Aulosira etc
Paspalum-notatum gras have loose symbiosis
(associative symbiosis) with Azotobacter paspali in
roots and Azospirillum with wheat, malze, sorghum
etc. Usually Azospirillum fix the atmospheric
nitrogen in free living condition.

Both Rhizobium & Frankia live freely in the soil but

fix nitrogen only when in symbiotic association with
host plant.
Wnzyme nitrogenase catalyses the conversion of
atmospheric N2 to NH3. It posses two units unit- Ist
is Mo-Fe protein & unit-IInd is Fe-S protein.

Nitrogenase is extremely sensitive to oxygen. So

to protect it from oxygen, leguminous nodules
contains on O2 scavanger called leghaemoglobin
(Lhb) which combines with O2 to form
oxyleghaemoglobin (LhbO2)
Legaemoglobin is pink or red in colour (globin part
synthesised by plant and haem part synthesised by
bacteria).

Many cyanobacteria capable of fixing nitrogen ar

fillamentous and contain thick walled cells called
heterocyst. These are the sites of nitrogen fixation.
Heterocyst lacks oxygen evolving photosystem II
thus do not evolve O2 and protect nitrogenase.
Biological N2 fixation involves contribution of
following genes :
(i) NOD gene of host plant
(ii) NOD, fix & NIF(Nitrogenase Including Factor)
genes of nitrogen fixing microbe.
Mechanism of Biological N2 fixation :

The atmospheric N2 is reduced by the addition of

hydrogen atoms.

The three bonds two nitrogen atom N = N or dinitrogen

are broken step by step & ammonia (NH3) is formed by
reduction of N º N and then ionisation of ammonia
(NH3)to form ammonium ions (NH4–1).
Ferredoxin helps in electron transfer.
N2 fixation requiers 3 components :
(i) A strong reducing agent – NADPH2 / FADH2 / NADH2
– from photosynthesis & respiration.

(ii) ATP-from respiration. (In symbiotic fixation from the

respiration of host cells).
Rhizobium – Legume symbiosis –
Rhizobium is a free-living, gram negative
nonsporulating, aerobic and motile rod shaped
bacterium. Rhizobia are more prominent in the
rhizosphere of leguminous plants.
Nodule formation - Nodule formation involves a
sequence of multiple interaction between Rhizobium
and roots of the host plant. Principal stages in the
nodule formation are summarised as follows :
(i) Rhizobia multiply and colonise the surroundings
of roots and get attached to epidermal and root
hair cells.
(ii) The root-hairs curl and the bacteria invade the
root – hair.
(iii) An infection thread is produced carrying the
bacteria into the cortex of the root, where they initiate
the nodule formation in the cortex and pericycle of
the root.
(iv) Then the bacteria are released from the thread
into the cells which leads to the differentiation of
specialised nitrogen fixing cells.
(v) The nodule thus formed, establishes a direct
vascular connection with the host for exchange of
nutrients.
UPTAKE AND ASSIMILATION OF NITROGEN
BY PLANTS (FATE OF AMMONIA)

Reductive amination – Inorganic NH 3 abrorbed

soil as NH4+ reacts with a-ketoglutaric acid to form
an amino acid glutamic acid. This process known
as Reductive amination or Amino acid
Biosynthesis.
Transmination – Transfer of Amino group from
glutamic acid to other keto acid is known as
transmination. This is process of formation of other
amino aicds from glutamic acid in plants, catalysed
by transaminase enzyme.
Glutamic acid + OAA  Aspartic acid + a-keto
glutaric acid
Glutamic acid + Pyruvic acid  Alanine + a-
ketoglutaric acid
Transportation of Assimilated N2 :
In plants transportation of assimilated N2 through
xylem occurs mainly in form of amides (Glutamine
and Asparagine), especially in leguminous plants.

Amides are more stable than amino acids and

posses high nitrogen to carbon ratio (2N to 5C in
glutamine, while glutamic acid posses 1N to 5C).
Formation of amides from amino acids by the
addition of amino group. (The hydroxyl part of acid
replaced by NH2 radicle) catalysed by enzyme and
is called Catalytic amidation.

In Soyabean, nitrogen is translocated as Ureides.

Now, this transported nitrogen is used in amino
acid synthesis and inturn amino acids are used in
protein synthesis and by herbivory, this nitrogen is
now transferred to the animals from plants.
Ammonification
After the death of plants and animals the protein
(organic nitrogen) present in dead biomass,
degraded or decomposed by some bacteria, in the
form of ammonia, the process is called
ammonification & the soil bacteria used in process
are called ammonifying bacteria.
e.g., Bacillus vulgaris, Bacillus ramosus, Bacillus
mycoides.
The ammonia produced by ammonification is
divided into three fractions –
(i) Some amount volatilises
(ii) Some amount converted into NH4+ ions
(iii) Most of the amount is converted into nitrate in
soil (Nitrification).
Nitrification
Oxidation of ammonia into nitrates by nitrifying bacteria
(Chemoautotrophs) is called nitrification.
Nitrate reduction
The nitrate thus formed is absorbed by plants and
transported to the leaves.
In leaves this nitrate is reduced to form ammonia
that finally forms the amine groups of amino acid.
NO3– ® NO2– - NH3
The process of nitrate reduction into ammonia is
called assimilatory nitrate reduction. It is catalysed
by enzyme nitrate reductase and nitrite reductase.
Denitrification
Some amount of nitrate present in the soil is also
reduced to nitrogen (N2) by the process of
Denitrification, this process is also called
deassimilatory nitrate reduction.
NO3– - NO2– - N2
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