TAMIL NADU AGRICULTURAL UNIVERSITY

CRP101
PRINCIPLES OF CROP PHYSIOLOGY
(2 +1)

LECTURE NOTES

Contents
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lecture title
an introduction and its importance in agriculture
role and significance of water diffusion, imbibition, osmosis and its significance, plasmolysis
Definition - field capacity, available soil water and permanent wilting point
Absorption of water - mode of water absorption active and passive absorption and factors affecting
absorption
Translocation of solutes - phloem and xylem transport
Transpiration - types - stewards theory of mechanism - significance, factors affecting transpiration and
guttation - antitranspirants
Mineral nutrition - introduction - criteria of essentiality of elements - macro, secondary and
micronutrients - soil less culture - sand and hydroponics
Mechanism of uptake - physiological role of nutrients
Foliar diagnosis - nutritional and physiological disorders - foliar nutrition.
Photosynthesis - requirements of photosynthesis - light, co2, pigments and H20
Mechanism of photosynthesis - light reaction - cyclic and non cyclic photophosphorilation - red drop emerson enhancement effect
Photosynthetic pathways - c3, c4 and cam
Differences between c3, c4 and cam pathways - factors affecting photosynthesis
Photorespiration - photorespiration process and significance of photorespiration
Respiration - glycolysis, tca and pentose phosphate pathway
Oxidative phosphorylation differences between oxidative phosphorylation and photophosphorylation.
respiratory quotient and energy budgeting in respiration.
Mid semester examination
Factors affecting respiration - difference
between photorespiration and dark respiration - role of
respiration
Protein and fat synthesis
Photoperiodism - short day, long day and day neutral plants phytochrome - role of phytochrome in
flowering and regulation of flowering
transmission of stimulus - theories of flowering
Vernalisation - mechanism of vernalisation and its significance - devernalisation
Source sink relationship - yield components - harvest index and its importance
Growth - growth curve, phases of growth and factors influencing growth
Growth analysis - LAI, LAD, SLW, SLA, LAR, NAR, RGR and CGR in relation to crop productivity
Plant growth regulators - growth hormones - definition and classification - physiological role of auxins
and GA.
Physiological role of cytokinin, ethylene and ABA - synthetic growth regulators and their uses in crop
productivity
Practical application of plant growth regulators in crop productivity
Environmental stresses - water stress - physiological changes - adaptation to drought and amelioration.
Temperature stress - physiological changes - low and high temperature - chilling injury - tolerance alleviation.
Low light and uv radiation stresses - salt stress - physiological changes and alleviation.
Global warming - physiological effects on crop productivity
Seed germination - physiological changes during seed germination
Abscission - senescence ripening types causes - physiological and biochemical changes and
regulation

Page
No

Lecture No: 1
AN INTRODUCTION AND ITS IMPORTANCE IN AGRICULTURE
Crop physiology is concerned with the processes and functions of the crops at
cellular, sub-cellular and whole plant levels in response to environmental variables and
growth. In short, physiology is the study of functional aspects of crop plants.
The role of crop physiology in different aspects of agriculture is discussed here.
1. Mineral Nutrition
The detection of deficiencies and toxicities of particular mineral nutrient elements
have enabled us to make adequate soil amendments for better plant growth. Several
physiological disorders of crop plants such as wheat, rice, pulses and oil seeds have been
successfully corrected with the help of knowledge on physiology. For example,
application of zinc has corrected khaira disease of paddy. Similarly, knowledge of water
relations and mineral utilization has led to improved crop management and now it is
possible to grow plants at places where they never grew, by providing proper
physiological conditions. Studies on chelates and chelating agents have gone a long way
in making unavailable elements available at functional sites and controlling toxicities of
heavy metals.
2. Photoperiodism and Vernalization
Researches on photoperiodism and vernalization have made it possible to grow
certain plants and make them flower even in off seasons by suitably altering the
photoperiods and providing low temperature treatments.
3. Production Physiology
Carbohydrates are major produce of plants and highly valued to human beings.
Hence, for increasing its yield capacity, three aspects may be considered: i) production
(Photosynthesis) ii) storage (sink potential), and iii) control of distribution in plants, i.e.
directing the food material efficiently towards storage organ (translocation of solute).
Higher translocation capacity towards storage organs like seeds helps to produce higher
yields. In case of rice, the sink capacity of panicles, as well as, the size and longevity of
the photosynthetic system after anthesis (flowering) can influence grain yield appreciably.
4. Photosynthesis
Green plants utilise less than one per cent of solar energy for the production of
food, and there are two types of plants, C3 and C4 based on CO2 assimilation. C3 plants re
less efficient photosynthesisers than C4 plants, though these plants are more precious to
human beings. C3 plants include pulses, oilseed crops, fibre crops, important cereals such
as wheat, rice, barley etc. Their lower efficiency is due to occurrence of photorespiration
which deviate Rubisco enzyme from photosynthesis. Therefore, there is a great need to
3

reduce wasteful process of photorespiration or to find out newer crop varieties with low
rate of photorespiration.
5. Plant Growth Physiology
Synthetic auxins and related compounds are being used for thinning of crops,
prevention of premature fruit drop, promotion of plant growth and yield, induction of
seedless or parthenocarpic fruits, promotion of root formation in cuttings for vegetative
propagation, budding or sprouting, induction of flowering, control of fruit set and quality,
hastening maturity, inducing dormancy in potato, controlling weeds etc.
Gibberellins have found great use in breaking dormancy, inducing uniform crop
emergence, producing staminate flowers in cucurbits, loosening fruit clusters, increasing
fruit size, hastening maturity, improving fruit quality, production of seedless ness,
increasing sugar content in sugarcane, inducing flowering etc.
Cytokinins have been widely used for increasing shelf life of fruits, quickening root
induction and producing efficient root system, increasing yield and oil contents of
groundnut, breaking dormancy, delaying senescence of living organisms, causing cell
division etc.
Ethylene has shown great potentials in making chemical harvesting possible,
thinning by causing abscission, inducing bulbing in onion and tillering in other crops,
causing dwarfness of plants, preventing lodging and inducing femaleness.
A number of other chemicals are also being used for causing male sterility,
overcoming incompatibility, environmental engineering and land maintenance. In welldeveloped countries, some growth regulators are frequently used in agriculture. For
example, chlormequat (CCC) is used as dwarfing agent in wheat. Ethephon is used to
induce flowering in pineapple and as sugarcane ripener. Maleic hydrazide is used as
growth retardant, for sucker control on tobacco, and as turf grass growth inhibitor.
Daminozide is used to enhance size and colour of various fruits. Glyphosine is used as
sugarcane ripener. Mepiquat chloride is a recent addition in the group of growth
retardants, having potential use in cotton, ground nut, banana and many vegetable crops.
Role of various hormones like, auxins, gibberellins, cytokinins and ethrel in
inducing and promoting flowering is now well documented. Ethrel is widely used to
increase the number of female flowers followed by higher yield in cucumber.
In several plants such as mango, apple, cotton etc., the fruits abscise and fall
before attaining maturity. Auxins have been found successful in preventing immature fall
of fruits and thus saving the enormous loss. Planofix (a formulation of NAA) is
commonly used in cotton, mango and coconut.
Tissue culture is the technique used to make successful in vitro growing of plant
4

parts under controlled aseptic conditions. Tissue culture practices are now widely used
and found immensely valuable in crop improvement programmes and hybridization
programmes. This technique is commonly used in:
*

Micro propagation in orchids, bananas, potatoes etc.

Production of disease free plants in potato, cassava, sugarcane, sweet potato etc.

Androgenic haploid and their use in breeding

Embryo rescue for successful hybridization, as in case of interspecific

hybridization between Phaseolus vulgaris and P. angustissimus.

Induction and selection of mutants

Somoclonal variations, as in case of wheat, potato and tomato

Protoplast technology, as in production of pomato (a hybrid of potato and tomato

produced by protoplasmic fusion)

6. Environment Physiology
The environmental factors influencing growth and yield of crops include
temperature, solar radiation (light), atmospheric carbon dioxide, water supply, air
humidity, wind velocity etc.
i) Temperature
Extreme temperatures are destructive to plant growth. The critically low and high
temperatures normally vary from one stage to another. The critical temperature will be
below 20C and above 30C for many of the crops. These temperatures also differ
according to variety, duration of critical temperature, diurnal changes and physiological
status of the plant. Temperature greatly influences the growth rate just after germination.
Within a temperature range of 22-31C, the growth rate increases almost linearly with
increasing temperatures.
Depending on the growth stages, injury to crop due to low temperature occurs
when the daily mean temperature drops below 20C. Common cool injuries are failure to
germinate, delayed seedling emergence, stunting, leaf discolouration, panicle tip
degeneration, incomplete panicle exertion, delayed flowering, high spikelet sterility and
irregular maturity. On the other hand, high temperatures cause high percentages of
spikelet sterility, when the temperatures exceed 35C at anthesis for more than 1 hour.
Vernalization: This is a method of inducing early flowering in plants by pre-treatment of
their seeds at very low temperatures. Practical utility of vernalization are i) crops can be
produced earlier ii) crops can be grown in the regions where they do not naturally
reproduce and iii) plant breeding work can be accelerated.
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ii) Solar Radiation

Most of radiant energy from the sun has a wavelength between 300 and 3000nm,
often referred as short wave radiation. Photosynthesis in green leaves uses solar energy in
wave lengths from 0.4 to 0.7 m (400-700 nm), often refer as Photosynthetically Active
Radiation (PAR). The solar radiation requirements of a crop differ from one growth stage
to another. For example, in rice, shading during the vegetative stage only, slightly affects
yield and yield components. Shading during the reproductive stage, however, has a
pronounced effect on spikelet number. During ripening, it reduces grain yield
considerably because of a decrease in the percentage of filled spikelets.
Flowering
The physiological mechanism responsible for flowering is found to be controlled by
duration of light (photoperiodism) and temperature (vernalization).
Photoperiodism: On the basis of length of photoperiod requirement the plants are
classified into: short-day plants, long-day plants and day-neutral plants.
Short-day plants: For flowering of short-day plants, the day length must not exceed a
certain critical value; the day length required is less than the critical length. Short-day
plants will not flower even if a flash of light is provided during the continuous dark
period. eg. Rice, onion, upland cotton and strawberry.
Long-day plants: Long-day plants require a photoperiod of more than a critical length,
which may vary from 14 to 18 hours. The best flowering of long day plants usually
occurs in continuous light. eg. Lettuce, radish, alfalfa, sugar beet, spinach etc.

Day-neutral plants: Their flowering is not affected by the length of the day. They can
flower even if the light provided is from few hours to continuous illumination. eg.
tomato, cucumber, cotton, pea, maize, sun-flower etc.
iii) CO2 concentration
Being one of the raw materials for photosynthesis, CO 2 concentration affects the
rate of photosynthesis markedly. Because of its very low concentration in the atmosphere
(360 ppm), it acts as a limiting factor for natural photosynthesis. The rate of
photosynthesis increases markedly with increase in the CO 2 level up to a certain extent.
Under full sunlight, photosynthesis increases up to 1000 ppm CO2.
iv) Water Supply
Water stress at any growth stage may reduce yield. The most common symptoms
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of water deficit are leaf rolling, leaf scorching, and impaired tillering, stunting and
delayed flowering, spikelet sterility and incomplete grain filling. Depending on
topography and rainfall patterns, low-lying areas may be subjected to different water
depths and to different duration. When a crop is submerged for a long time during critical
growth stages, the grain yield is reduced.

Lecture No: 2

Role and significance of water

Diffusion, imbibition, osmosis and its significance,
plasmolysis.
Water is said to be the liquid of life. Because life originated in organs,
environmental and in the course of evolution it became fully dependent upon water in a
number of ways.
Importance of water to plants
1. Water is the main constituent of protoplasm comprising up to about 90-95 per cent
of its total weight. In the absence of water, protoplasm becomes inactive and is
oven killed.
2. Different organic constituents of plants such as carbohydrates proteins, nucleic
acid and enzymes etc. Lose their physical and chemical properties in the absence
of water.
3. Water participates directly in many metabolic processes. Inter conversion of
carbohydrates and organic acids depend upon hydrolysis and condensation
reaction.
4. Water increases the rate of respiration. Seeds respire fast in the presence of water.
5. Water is the source of hydrogen atom for the reduction of CO 2 in the reaction of
photosynthesis.
6. Water acts as a solvent and acts as a carrier for many substance. If forms the
medium in which several reactions take place.
7. Water present in the vacuoles helps in maintaining the turgidity of the cells which
is a must for proper activities of life and to maintain this from and structure.
8. Water helps in translocation of solutes
9. In tropical plants, water plays a very important role of thermal regulation against
high temperature.
10. The elongation phase of cell growth depends on absorption of water.
Diffusion, osmosis and imbibition
The movement of materials in and outt of the cells in plants taken place in a
solution or gaseous form. Although the exert process of this is not very clear, three
physical process are usually involved in it. They are diffusion, osmosis and imbibition.
The movement of particles or molecules from a region of higher concentrations to
a region of lower concentration is called as diffusion. The rate of diffusion of gases is
faster than liquids or solutes. The diffusion particles have a certain pressure called as the
diffusion pressure which is directly proportional to the number as concentration of the
diffusing particles. These forms the diffusion takes place always from a region of higher
diffusion pressure to a region of lower diffusion pressure (i.e) along a diffusion pressure
gradient. The rate of diffusion increases if,

i)
ii)
iii)
iv)

Diffusion pressure gradient is steeper

Temperature is increased
Density of the differing particles is lesser
Medium through which diffusion occurs is less concentrated.

Diffusion of more than one substance at the same time and place may be at
different rates and in different direction, but is independent of each other. A very
common example of this is the gaseous exchange in plants.
Beside osmotic diffusion the above mentioned simple diffusion also plays a very
important role in the life of the plants.
-

It is an essential step in the exchange of gases during respiration and

photosynthesis
During passive salt uptake, the ions are absorbed by diffusion
It is important in stomatal transpiration as the last step in the pollen, where
diffusion of water vapour from the interrelation space into the outer atmosphere
occurs through open stomata.

Osmosis
The diffusion of solvent molecules into the solution through a semi permeable
membrane is called as osmosis (some times called as Osmotic diffusion). In case there
are two solutions of different concentration separated by the semi permeable membrane,
the diffusion of solvent will take place from the less concentrated suitable into the more
concentrated solution till both the solutions attain equal concentration.

Osmotic pressure
As a result of the separation of solution from its solvent (or) the two solutions by
the semi permeable membrane, a pressure is developed in solution to the pressure by
dissolved solutes in it. This is called as osmotic pressure (O.P). OP is measured interms
of atmospheres and is directly proportional to the concentration of dissolved solutes in
the solution. More concentration solution has higher O.P. O.P of a solution is always
higher than its pure solvent.
During osmosis, the movement of solvent molecules taken place form the solution
whose osmotic pressure is lower (i.e less concentration as hypotonic) into the solution
whose osmotic pressure is higher (i.e, more concentrated as hypertonic).
Osmotic diffusion of solvent molecules will not take place if the two solutions
separated by the semi permeable membrane are of equal concentration having equal

Osmotic pressures (i.e., they are isotonic). In plant cells, plasma membrane and
tonoplant act as selectively permeable or differentially permeable membrane.

Endo osmosis
Of a living plant cell is placed in water or hypotonic solution whose O.P is lower
than cell sap, water interms into the cell sap by osmosis and the process is called end
osmosis. As a result of entry of water with the cell sap, a pressure is developed which
press the protoplasm against the cell wall and become turgid. This pressure is called a
turgor pressure.
Consequence of the turgor pressure is the wall pressure which is exerted by the
elastic cell wall against the expanding protoplasm. At a given time, turgor pressure (T.P)
equals the wall pressure (W.P).
T.P = W.P
Exosmosis
If on the other hand, the plant cell is placed in hypertonic solution (whose O.P is
higher than cell sap) the water cover out the cell sap into the outer solution and the cell
becomes flaccid. This process is known as exosmosis. Cell (or) tissue will remain as
such in isotonic solution.

Significance of osmosis in plants

1.
2.
3.
4.

Large quantities of water are absorbed by roots from the soil by osmosis
Cell to cell movement of waster and other substances dissolve is involves osmosis
Opening and closing of stomata depend upon the turgor pressure of guard cells
Due to osmosis, the turgidity of the cells and hence the shape or from of them
organs is maintained.
5. The resistance of plants to drought and frost increases with increase in osmotic
pressure to later cells
6. Turgidity of the cells of the young seedling allows them to come out of the soil.

Imbibition
Certain substances if placed in a particular liquid absorb it and swell up. For
example, when some pieces of grass or dry wood or dry seeds are placed in water they
absorb the water quickly and swell up considerably so that their volume is increased.
These substances are called as imbibants and the phenomenon as imbibition, certain force

10

of attraction is existing between imbibants and the involved substance. In plants, the
hydrophilic colloids viz., protein and carbohydrates such as starch, cellulose and pectic
substance have strong altercation towards water.
Imbibition plays a very important role in the life of plants. The first step in the
absorption of water by the roots of higher plants is the imbibition of water by the cell
walls of the root hairs. Dry seeds require water by imbibition for germination.
As a result of imbibition, a pressure is developed which is called as imbibition
pressure or matric potential (m). It is analogous to the osmotic potential of a solution.
With reference to pure water, the values of m are always negative. The water potential
of an imbibant is equal to its matric potential plus any turgor or other pressure (pressure
potential) which may be imposed upon the imbibant.
w = m + P
If the imbibant is unconfined to turgor or such pressure, the equation can be
significant to
w =m
Plasmolysis
When a plant cell or tissue is placed in a hypertonic solution water cover out from
the cell sap into the outer solution of exosmosis and the protoplasm begins to sprinkler or
contract. The protoplasm separate from the cell wall and assures a spherical form and
them phenomenon is called plasmolysis. Incipient plasmolysis is stage where protoplasm
begins to contract from the cell wall. If a plasmolysed cell in tissue is placed in water, the
process of endosmosis take place. Water enters into the cell sap, the cell becomes turgid
and the protoplasm again assumes it normal shape and position. This phenomenon is
called deplasmolysis.

Advantages of plasmolysis
1. It indicates the semi permeable nature of the plasma membrane.
2. It is used in determine the osmotic pressure of the cell sap.

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3. Plasmolysis is used in salting of meat and fishes. Addition of concentrated sugar

solution to jam and jellies check the growth of fungi and bacteria which become
plasmolysed in concentrated solution.

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Lecture No: 3

Definition - field capacity, Available soil water

and permanent wilting point
Field capacity or water holding capacity of the soil
After heavy rain fall or irrigation of the soil some water is drained off along the
slopes while the rest percolates down in the soil. Out of this water, some amount of water
gradually reaches the water table under the force of gravity (gravitational water) while the
rest is retained by the soil. This amount of water retained by the soil is called as field
capacity or water holding capacity of the soil.
Field capacity is affected by soil profiles soil structure and temperature.
The effective depth of a soil, as determined by physical and chemical barriers, together
with the clay content of the soil within that depth, determine the water holding capacity
of the profile, and how much of the water is available to plants. Effective soil depth
varies between plant species. Wheat is used as the benchmark plant in this assessment.
Available water holding capacity rankings are estimated from soil texture, structure and
stone content within the potential root zone of a wheat plant.
Water-holding capacity is controlled primarily by soil texture and organic matter.
Soils with smaller particles (silt and clay) have a larger surface area than those with larger
sand particles, and a large surface area allows a soil to hold more water. In other words, a
soil with a high percentage of silt and clay particles, which describes fine soil, has a
higher water-holding capacity. The table illustrates water-holding-capacity differences as
influenced by texture. Organic matter percentage also influences water-holding capacity.
As the percentage increases, the water-holding capacity increases because of the affinity
organic matter has for water.

13

It is the water content of the soil after downward drainage of gravitational water.
It is the capillary capacity of a soil. It is the upper limit of soil water storage for the plant
growth. At field capacity, the soil water potential is 0.1 to 0.3 bars.

Water potentials
Every component of a system possesses free energy capable of doing work under
constant temperature conditions.

For non-electrolytes, free energy / mole is known as

chemical potential. With refuse to water, the chemical potential of water is called as
water potential. The chemical potential is denoted by a Greek letter Psi ().
For pure water, the water potential is O. The presence of solute particles will
reduce the free energy of water or decrease the water potential. Therefore it is expressed
in vegetative value.
It is therefore, water potential of solution is always less than O so in negative
value.
For solutions, water potential is determined by three internal factors i.e.
w

= w + s + p

= is the solute potential or osmotic potential

= pressure potential or turgor potential

= is the matric potential. Matric potential can be measured for the water
molecules adhering on the soil particles and cell wall.

In plant system, the matric potential is disregarded.

Therefore,
w = s + p

Osmotic pressure
Osmotic pressure is equivalent to osmotic potential but opposite in sign.

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Osmotic pressure in a solution results due to the presence of solutes and the
solutes lower the water potential. Therefore osmotic pressure is a quantitative index of
the lowering of water potential in a solution and using thermodynamic terminology is
called as osmotic potential.
Osmotic pressure and osmotic potential are numerically equal but opposite in
sign.
Osmotic pressure has positive sign
Osmotic potential has negative sign (s)
For eg.
IA OP = 20 atm.
w

= - 20 atm

Turgor pressure
In plant cell, the turgor pressure results due to the presence of water molecules is
turgor pressure. The potential created by such pressures is called presume potential (p)
In a normal plant cell, the water potential
w = s + p partially turgid cell
(High)
w = Zero

- Fully turgid cell

(Highest)
w = s

- Flaccid cell or plasmolysed cell

(Lowest)

Water relation
Water form the major constituent of living (cells) things and the cells originated in
a highly aqueous medium and all the vital processes of the life are carried out in it.

15

Besides, water predominately arts as a source of hydrogen to plants and is released by the
photolysis of water during photosynthesis.
In living tissue, water is the medium for many biochemical reactions and
extraction process.

Inorganic nutrients, photosynthesis, bases and hormones are all

transported in aqueous solution. Evaporation of water can control the temperature of leaf
on canopy soil nutrients are available to plant roots only when dissolved in water. In
short, water is essential for life and plays a unique role in virtually all biological process.
Example:
There are 2 cells A and B in contact with each other, cell A has a pressure
potential (turgor pressure) of 4 bars and certain sap with an osmotic potential of -12 bars.
Cell B has presume potential of 2 bars and certain sap with osmotic potential of -5
bars.
Then,
w of cell A

= s + p
= -12 + (+4)
= -8 bars

w of cell B

= -5+(+2)
-3 bars

Hence, water will move from cell B to cell A (i.e., towards lower or more negative
water potential) with a form of (-8-(-3) = -5 bars.
Diffusion Pressure Dficit (DPD)
(Suction pressure)
Diffusion pressure of a solution is always lower than its pure solvent. The
difference between the diffusion pressure of the solution and its solvent at a particular
temperate and atmosphere conditions is called as diffusion pressure deficit (D.P.D). If the
solution is more concentrated D.P.D increases but it decreases with the dilution of the
solution,
16

D.P.D of the cell sap or the cells is a measure of the ability of the cells to absorb
water and hence is often called as the suction pressure (S.P). It is related with osmotic
pressure (O.P) and turgor pressure (T.P) of cell sap and also the wall pressure (W.P) as
follows.
D.P.D. (S.P) = O.P W.P
But
(W.P) = T.P
D.P.D = O.P T.P
Due to the entry of the water the osmotic pressure of the cell sap decreases while
its turgor pressure is increased so much so that in a fully turgid cell T.P equals the O.P
O.P = T.P

= D.P.D = O

In fully plasmolysed cells: T.P = O

So

D.P.D = O.P

D.P.D. incase of plant cells is not directly proportional to their osmotic pressure or
the concentration of the cell sap but depend both on O.P and T.P. Higher osmotic
pressure of the cell sap is usually accompanied by lower turgor pressure so that its D.P.D
is greater and water enters into it. But sometimes it is possible that two cells are in
contact with each other one having higher O.P and also higher turgor pressure than the
other cell and still its does not draw water. It is because of its lower D.P.D., no matter is
O.P is higher.
Cell a

Cell b

O.P

= 25 atm.

O.P

= 30 atm

T.P

= 15 atm.

T.P

=10 atm.

D.P.D = 10 atm.

D.P.D = 30 atm.

Cell a

Cell b

17

O.P

= 35 atm.

O.P

= 40 atm

T.P

= 10 atm.

T.P

= 20 atm.

D.P.D = 25 atm.

D.P.D = 20 atm.

Entry of water into the cell depends on D.P.D and not on O.P only

18

Lecture No: 4

Absorption of water - mode of water absorption active

and passive absorption and factors affecting absorption
Mechanism of absorption of water
In higher plants water is absorbed through root hairs which are in contact with soil
water and form a root hair zone a little behind the root tips. Root hairs are tubular hair
like prolongations of the cells of the epidermal layer (when epidermis bears root hairs it
is also known as pilloferous layer) of the roots. The walls of root hairs are permeable and
consist of pectic substances and cellulose which are strongly hydrophilic in nature root
hairs contain vacuoles filled with cell sap. When roots elongate, the older root hairs die
and new root hairs are developed so that they are in contact with fresh supplies of water
in the soil.

Mechanism of water absorption

1. Active absorption of water
In this process the root cells play active role in the absorption of water and
metabolic energy released through respiration is consumed active absorption may be of
two kinds.
(A) Osmotic absorption
Water is absorbed from the soil into the xylem of the roots according to osmotic
gradient.
(B) Non-osmotic absorption
Water is absorbed against the osmotic gradient.

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2. Passive absorption of water

It is mainly due to transpiration, the root cells do not play active role and remain
passive.
I. Active osmotic absorption of water
First step in osmotic the osmotic absorption of water is the imbibition of soil
water by the hydrophilic cell walls of root hairs. Osmotic pressure of the cell sap of root
hairs is usually higher than the OP of the soil water. Therefore, the DPD and suction
presume in the root hairs become higher and water from the cell walls enters into them
through plasma membrane by osmotic diffusion. As a result, OP, suction pressure and
DPD of root hairs how become lower, while their turgor pressure is increased.
Now the cortical cells adjacent to root hairs have high OP, SP & DPD in
comparison to the root hairs. Therefore, water is drawn into the adjacent cortical cells
from root hairs by osmotic diffusion. In the same way, by cell to cell osmotic diffusion
gradually reaches the inner most cortical cells and the endodermis. Osmotic diffusion of
water into endodermis takes place through special thin walled passage cells because the
other endodermis cells have casparian strips on thin walls which are impervious to water.
Water from endodermis cells is down into the cells of pericycle by osmotic
diffusion which now become turgid and their suction pressure in decreased.
In the last step, water is drawn into xylem from turgid pericycle cells (In roots the
vascular bundles are radical and protoxylem elements are in contact with pericycle). It is
because in the absence of turgor presume of the xylem vessels, the SP of xylem vessels
become higher than SP of the cells of the pericycle when water enters into xylem from
pericycle a presume is developed in the xylem of roots which can raise the water to a
certain height in the xylem. This pressure is called as root pressure.
(b) Active non-osmotic absorption of water
Sometimes, it has been observed that absorption of water takes place even when
OP of soil water is high than OP of cell sap. This type of absorption which is non-osmotic
20

and against the osmotic gradient requires the expenditure of metabolic energy probably
through respiration.

2. Passive absorption of water

Passive absorption of water takes place when rate of transpiration is usually high.
Rapid evaporation of water from the leaves during transpiration creates a tension in water
in the xylem of the leaves. This tension is transmitted to water in xylem of roots through
the xylem of stream and water rises upward to reach the transpiring surfaces. As the
results soil water enters into the cortical cells through root hairs to reach the xylem of
roots to maintain the supply of water. The force of this entry of water is created in leaves
due to rapid transpiration and hence, the root cells remain passive during this process.

External factors affecting absorption of water

1. Available soil water
Sufficient amount of water should be present in the soil in such form which can
easily be absorbed by the plants. Usually the plants absorb capillary water i.e water
present in films in between soil particles other forms of water in the soil eg. Hygroscopic
water, combined water, gravitational water etc. is not easily available to plants.
Increased amount of water in the soil beyond a certain limit results in poor
aeration of the soil which retards metabolic activities of root cells like respiration and
hence, the rate of water absorption is also retarded.
2.Concentration of soil solution
Increased concentration of soil solution (due to presence of more salts in the soil)
results in higher OP. If OP of soil solution will become higher than the OP of cell sap in
root cells, the water absorption particularly the osmotic absorption of water will be
greatly suppressed. Therefore, absorption of water is poor in alkaline soils and marshes.

21

3.Soil air
Absorption of water is retarded in poorly aerated soils because in such soils
deficiency of O2 and consequently the accumulation of CO2 will retard the metabolic
activities of roots like respiration. This also inhibits rapid growth and elongation of the
roots so that they are deprived of fresh supply of water in the soil. Water logged soils are
poorly aerated and hence, are physiologically dry. They are not good for absorption of
water.
4.Soil temperature
Increase in soil temperature up to about 30C favours water absorption. At higher
temperature water absorption is decreased. At low temperature also water absorption
decreased so much so that at about 0C, it is almost decreased. This is probably because
at low temperature.
1. The viscosity of water and protoplasm is increased
2. Permeability of cell membrane is decreased
3. Metabolic activity of root cells are decreased
4. Root growth and elongation of roots are checked.

22

Lecture No: 5
TRANSLOCATION OF SOLUTES - PHLOEM AND XYLEM
TRANSPORT
Translocation of organic solutes
The movement of organic food materials or the solutes in soluble form one place
to another in higher plants is called as
Translocation of organic solutes
Directions of translocation
Translocation of organic solutes may take place in the following directions.

1. Downward translocation
Mostly, the organic material is manufactured by leaves and translocated
downward to stem and roots for consumption and storage.

2. Upward translocation
It takes place mainly during the germination of seeds, tubers etc. When stored
food after being converted into soluble form is supplied to the upper growing part of the
young seedling till it has developed green leaves.
Upward translocation of solutes also takes place through stem to young leaves,
buds and flowers which are situated at the tip of the branch.

3. lateral translocation
Radical translocation of organic solutes also takes place in plants from the cells of
the pith to cortex.

23

Path of the translocation of organic solutes

1. Path of downward translocation
Downward translocation of the organic solutes takes place through phloem. This
can be proved by the ringing experiment.

2. Path of upward translocation

Although translocation of organic solutes take place through phloem, but under
certain conditions it may take place through xylem.

3. Path of lateral translocation

Lateral translocation from pith to cortex takes place through medullary
rays.

Mechanism of translocation
Various theories have been put forward to explain the mechanism of phloem
conduction. Among them Munchs (1930) hypothesis is mot convincing.

Munchs mass flow on pressure flow hypothesis

According to this hypothesis put forward by Much (1930) and others, the
translocation of organic solutes takes place though phloem along a gradient of turgor
pressure from the region of higher concentration of soluble solutes (supply end) to the
region of lower concentration (consumption end). The principle involved in this
hypothesis can be explained by a simple physical system as shown in Fig.

Two members X and Y permeable only to water and dipping in water are
connected by a tube T to form a closed system membrane X contains more concentrated
sugar solution than in membrane Y.
24

Due to higher osmotic presence of the concentrated sugar solution in the

membrane X, water enters into it so that its turgor pressure is increased. The increase in
turgor pressure results in mass flow of sugar solution to membrane Y though the T till the
concentration of sugar solution in both the membrane is equal.
In the above system it could be possible to maintain continuous supply of sugars
in membrane X and its utilization on conversion into insoluble form in membrane Y, the
flow of sugar solution from X to Y will continue indefinitely.
According to this theory, a similar analogous system for the translocation of
organic solutes exists in plants. As a result of photosynthesis, the mesophyll cells in the
leaves contain high concentration of organic food material in them in soluble form and
correspond to membrane X or supply end.
The cells of stem and roots where the food material is utilized or converted into
insoluble form correspond to membrane Y or consumption end. While the sieve tubes in
phloem which are placed and to end correspond to the tube T.
Mesophyll cells draw water from the xylem of the leaf due to higher osmotic
pressure and suction presume of their sap so that their turgor pressure is increased. The
turgor presume in the cells of stem and the roots is comparatively low and hence, the
soluble organic solutes begin to flow en mass from mesophyll through phloem down to
the cells of stem and the roots under the gradient of turgor presume. In the stem and the
roots, the organic solutes are either consumed or converted into insoluble form and the
excess water is released into xylem through cambium.

XYLEM TRANSPORT
ASCENT OF SAP
The water after being absorbed by the roots is distributed to all parts of the plants.
In order to reach the topmost part of the plant, the water has to move upward through the
stem. The upward movement of water is called as Ascent of sap.
25

Ascent of sap can be studied under the following two headings.

1. Path of ascent of sap
2. Mechanism of ascent of sap.

1. Path of ascent of sap

Ascent of sap takes place through xylem. It can be shown by the experiment.
0

A leafy twig of Balsam plant (it has semi transpiration stem) is cut under water (to

avoid entry of air bubble through the cut end of the stem) and placed in a beaker
containing water with some Eosine (a dye) dissolved in it.
1

After sometimes coloured lines will be seen moving upward in the stem. If

sections of stem are cut at this time, only the xylem elements will appear to be filled with
coloured water.
2. Ringing experiment
A leafy twig from a tree is cut under water and placed in a beaker filled with
water. A ring of bark is removed from the stem. After sometime it is observed that the
leaves above the ringing part of the stem remain fresh and green. It is because water is
being continuously supplied to the upper part of the twig through xylem.
B. Mechanism of ascent of sap
In small trees and herbaceous plants, the ascent of sap can be explained easily, but
in tall trees like Eucalyptus and conifers reaching a height of 300-400 feet), where water
has to rise up to the height of several hundred feet, the ascent of sap, it feet, becomes a
problem. To explain the mechanism of Ascent of sap, a number of theories have been put
forward.
a. vital theory
b. root pressure theory
c. physical force theory
d. transpiration pull and cohesion of water theory
26

A. Vital theories
According to vital theories, the ascent of sap is under the control of vital activities
in the stem.
1. According to Godlewski (1884) Ascent of sap takes place due to the pumping
activity xylem tissues which are living.
2. According to Bose (1923) upward translocation of water takes place due to
pulsatory activity of the living cells of the inner must cortical layer just outside
the endodermis.

B. Root pressure theory

Although, root pressure which is developed in the xylem of the roots can raise
water to a certain height but does not seem to be an effective force in ascent of sap due to
the following reasons. Magnitude of root pressure is very low (about 2 atmos).
0

Even in the absence of root pressure, ascent of sap continues. For example, when

leafy twig is cut under water and placed in a beaker full of water it remains fresh and
green for sufficient long time.

C. Physical force theories

Various physical forces may be involved in A. of sap.

1. Atmospheric pressure
This does not seem to be convincing because
0

it cannot act on water present in xylem in roots

Incase it is working, and then also it will not be able to raise water beyond 34.

27

2. Imbibition
Sachs (1878) supported the view that ascent of sap could take place by imbibition
through the walls of xylem. But imbibitional force is insignificant in the A. of sap
because it takes place through the lumen of xylem elements and not through walls.

3. Capillary force
In plants the xylem vessels are placed one above the other forming a sort of
continuous channel which can be compared with long capillary tubes and it was thought
that as water rises in capillary tube due to capillary force in the same manner ascent of
sap takes place in the xylem.

D. Transpiration pull and cohesion of water theory

This theory was originally proposed by Dixon and Jolly (1894) later supported
and elaborated by Dixon (1924). This theory is very convincing and has now been
widely supported by many workers.
Although H- bond is very weak (Containing about 5 K cal energy) but they are
present in enormous numbers as incase of water, a very strong mutual force of attraction
or cohesive force develops between water molecules and hence they remain in the form
of a continuous water column in the xylem. The magnitude of this force is very high (up
to 350 atm), therefore the continuous water column in the xylem cannot be broken easily
due to the force of gravity or other abstractions offered by the internal tissues in the
upward movement of water.
The adhesive properties of water i.e. attractions between the water molecules and
the containers walls (here the walls of xylem) further ensure the continuity of water
column in xylem.
When transpiration takes place in the leaves at the upper parts of the plant, water
evaporates from the intercellular spaces of the leaves to the outer atmosphere through
28

stomata. More water is released into the intercellular spaces from mesophyll cells. In
turn, the mesophyll cells draw water from the xylem of the leaf. Due to all this, a tension
is created in the xylem elements of the leaves. This tension is transmitted downward to
water in xylem elements of the root through the xylem of petiole and stem and the water
is pulled upward in the form of continuous unbroken water column to reach the
transpiring surfaces up to the top of the plant

29

Lecture No: 6
TRANSPIRATION - TYPES - STEWARDS THEORY OF MECHANISM SIGNIFICANCE, FACTORS AFFECTING TRANSPIRATION AND
GUTTATION - ANTITRANSPIRANTS
Although large quantities of water are absorbed by plant from the soil but only a
small amount of it is utilized. The excess of water is lost from the aerial parts of plants in
the form of water vapours. This is called as transpiration.

Transpiration is of 3 types
1. Stomatal transpiration
Most of the transpiration takes place through stomata.

Stomata are usually

confined in more numbers on the lower sides of the leaves. In monocots. Eg. Grasses
they are equally distributed on both sides. While in aquatic plants with floating leaves
they are present on the upper surface.

2. Cuticular transpiration
Cuticle is impervious to water, even though, some water may be lost through it. It
may contribute a maximum of about 10% of the total transpiration.

3. Lenticular transpiration
Some water may be lost by woody stems through lenticells which is called as
lenticular transpiration.

30

Mechanism of stomatal transpiration

The mechanism of stomatal transpiration which takes place during the day time
can be studied in 3 steps.
1

Osmotic diffusion of water in the leaf from xylem to intercellular space above the

stomata through the mesophyll cells.

2

Opening and closing of stomata (stomatal movement)

Simple diffusion of water vapours from intercellular spaces to other atmosphere

through stomata.

Inside the leaf the mesophyll cells are in contact

With xylem, and on the other hand with intercellular space above the stomata
When mesophyll cells draw water from the xylem they become turgid and their
diffusion pressure deficit (DPD) and osmotic pressure (OP) decreases with the
result that they release water in the form of vapour in intercellular spaces close to
stomata by osmotic diffusion. Now in turn, the O.P and D.P.D of mesophyll cells
become higher and hence, they draw water form xylem by osmotic diffusion.

3. Opening and closing of stomata (Stomatal movement)

The stomata are easily recognized from the surrounding epidermal cells by their
peculiar shape. The epidermal cells that immediately surround the stomata may be
similar to other epidermal cells or may be different and specialized. In the latter case,
they are called as subsidiary cells.
The guard cells differ from other epidermal cells also in containing chloroplasts
and peculiar thickening on their adjacent surface (in closed stomata) or on surfaces.
Consequent to an increase in the osmotic pressure (OP) and diffusion pressure
deficit (DPD) of the guard cells (which is due to accumulation of osmotically active
substances), osmotic diffusion of water from surrounding epidermal cells and mesophyll
31

cells into guard cells follows. This increase the turgor pressure (TP) of the guard cells
and they become turgid. The guard cells swell, increase in length and their adjacent
thickened surfaces starch forming a pore and thus the stomata open.
On the other hand, when OP and DPD of guard cells decrease (due to depletion of
osmotically active substances) relative to surrounding epidermal and mesophyll cells,
water is released back into the latter by osmotic diffusion and the guard cells become
flaccid. The thickened surfaces of the guard cells come close to each other, thereby
closing the stomatal pore and stomata.

Osmotic diffusion of water into guard cells occur when their osmotic pressure
increases and water potential decreases (i.e become more negative) related to those of
surrounding epidermal and mesophyll cells. The guard cells become flaccid when their

32

osmotic pressure decreases relative to the surrounding cells (Movement of water takes
place from a region of higher water potential to a region of lower water potential.
These may be several different agents or mechanisms which control stomatal
movements.
0

Hydrolysis of starch into sugars in guard cells

Synthesis of sugars or organic acids in them

The active pumping of K+ ions in the guard.

1. Hydrolysis of starch into sugars in guard cells

Starch sugar Inter conversion theory
This classical theory is based on the effect of pH on starch phosphorylase enzyme
which reversibly catalyses the conversion of starch + inorganic phosphate into glucose -1
phosphate.
During the day, pH is guard cells in high. This favours hydrolysis of starch
(which is insoluble into glucose -1- phosphate (which is soluble) so that osmotic pressure
is increased in guard cells.
Consequently water enters, into the guard cells by osmotic diffusion from the
surrounding epidermal and mesophyll cells. Guard cells become turgid and the stomata
open.
During dark, reverse process occurs. Glucose 1- phosphate is converted back into
starch in the guard cells thereby decreasing osmotic pressure. The guard cell release
water, become flaccid and stomata become closed.
Light high pH
Starch
(Insoluble)

+Pi

Glucose-1-phosphate
Dark low pH

(Soluble)

33

According to Steward 91964), the conversion of starch and inorganic phosphate

into glucose-1-phosphate does not cause any appreciable change in the osmotic pressure
because the inorganic phosphate and glucose-1-phosphate are equally active osmotically.
In this scheme he has suggested that,
0

Glucose-1-phosphate should be further converted into glucose and inorganic

phosphate for the opening of stomata.

1

Metabolic energy in the form of ATP would be required for the closing of stomata

which probably comes through respiration.

Starch
pH 5.0

pH 7.0

Glucose-1-phosphate
Hexokinase + ATP
O2 Resp.

Glucose-6-phosphate
Phosphatase
Glucose + Pi

2. Synthesis of sugars or organic acids in Guard cells

During day light photosynthesis occurs in guard cells as they contain chloroplast.
The soluble sugars formed in this process may contribute in increasing the osmotic
potential of guard cells and hence resulting in stomatal opening. However, very small
amounts of soluble sugars (osmotically active) have been extracted from the guard cells
which are insufficient to affect water potential.
As a result of photosynthesis CO2 concentration in guard cells decreases which
leads to increased pH up of organic acids, chiefly malic acid during this period in guard
cells. The formation of malic acid would produce proton that could operate in an ATPdriven proton K+ exchange pump moving protons into the adjacent epidermal cells and K

34

ions into guard cells and thus may contribute in increasing the osmotic pressure of the
guard cells and leading to stomatal opening.
Reverse process would occur in darkness.

3. ATP Driven proton (H+) K exchange pump mechanism in Guard cells

According to this mechanism, there is accumulation of K+ ions in the guard cells
during day light period. The protons (H+) are pumped out from the guard cells into the
adjacent epidermal cells and in exchange K+ ions are mediated through ATP and thus are
an active process.

ATP is generated in non-cyclic photophosphorylation in

photosynthesis in the guard cells. The ATP required in ion exchange process may also
come through respiration.
The accumulation of K ion is sufficient enough to significantly decrease the water
potential of guard cells during day light. Consequently, water enters into them from the
adjacent epidermal and mesophyll cells thereby increasing their turgor pressure and
opening the stomatal pore.
Reverse situation prevails during dark when stomata are closed. There is no
accumulation of K in g cells in dark.
(iii) The last step in the mechanism of transpiration is the simple diffusion of water
vapours from the intercellular spaces to the atmosphere through open stomata. This is
because the intercellular spaces are more saturated with moisture is comparison to the
outer atmosphere in the vicinity of stomata.

Significance of Transpiration
0

Plants waste much of their energy in absorbing large quantities of water and most

of which is ultimately lost through transpiration.

0

Some people thin that Transpiration as advantageous to plant.

Others regard it as an unavoidable process which is rather harmful.

35

Advances of transpiration
1. Role of movement of water
Plays an important role in upward movement of water i.e Ascent of sap in plants.

2. Role in absorption and translocation of mineral salts

Absorption of water and mineral salts are entirely independent process. Therefore
transpiration has nothing to do with the absorption of mineral salts.
However, once mineral salts have been absorbed by the plants, their further
translocation and distribution may be facilitated by transpiration through translocation of
water in the xylem elements.
3. Role of regulation of temperature
Some light energy absorbed by the leaves is utilized in photosynthesis; rest is
converted into heat energy which raises their temperature.

Transpiration plays an

important role in controlling the temperature of the plants. Rapid evaporation of water
from the aerial parts of the plant through transpiration brings down their temperature and
thus prevents them from excessive heating.
B. Transpiration as a necessary evil
1. When the rate of transpiration is high and soil is deficient in water, an internal water
deficit is created in the plants which may affect metabolic processes
2. Many xerophytes have to develop structural modification and adaptation to check
transpiration.
3. Deciduous tress has to shed their leaves during autumn to check loss of water.
But, in spite of the various disadvantages, the plants cannot avoid transpiration
due to their peculiar internal structure particularly those of leaves.

36

Their internal

structure although basically mean for gaseous exchange for respiration, P.S. etc. is such
that it cannot check the evaporation of water. Therefore, many workers like Curtis (1926)
have called transpiration as necessary evil.
Factors affecting transpiration rate
A. External factors
1. Atmospheric humidity
In humid atmosphere, (when relative humidity) is high), the rate of transpiration
decreases. It is because atmosphere is more saturated with moisture and retards the
diffusion of water vapour from the intercellular spaces of the leaves to the outer
atmosphere through stomata.
In dry atmosphere, the RH is low and the air is not saturated with moisture and
hence, the rate of transpiration increases.
2. Temperature
An increase in temperature brings about an increase in the rate of transpiration by
1. lowering the relative humidity
2. Opening of stomata widely
3. Wind
i. When wind is stagnant (not blowing), the rate of transpiration remains normal
ii. When the wind is blowing gently, the rate of transpiration increases because it removes
moisture from the vicinity of the transpiration parts of the plant thus facilitating the
diffusion of waster vapour from the intercellular spaces of the leaves to the outer
atmosphere though stomata.
iii. When the wind is blowing violently, the rate of transpiration decreased because it
creates hindrance in the outward diffusion of water vapours from the transpiring part and
it may also close the stomata.
4. Light
Light increases the rate of transpiration because
37

In light stomata open

It increases the temperature

In dark, due to closure of stomata, the stomatal transpiration is almost stopped.

5. Available soil water

Rate of transpiration will decrease if there is not enough water in the soil in such
from which can be easily absorbed by the roots.
6. CO2
An increase in CO2 concentration in the atmosphere (Ova the usual concentration)
more so inside the leaf, leads towards stomatal closure and hence it retards transpiration.
B. Internal factors
1. Internal water conditions
It is very essential for transpiration. Deficiency of water in the plants will result
in decrease of transpiration rate. Increase rate of transpiration containing for longer
periods often create internal water deficit in plants because absorption of water does not
keep pace with it.

2. Structural features
The number, size, position and the movement of stomata affect rate of
transpiration. In dark stomata are closed and stomatal transpiration is checked. Sunken
stomata help in reducing the rate of stomatal transpiration. In xerophytes the leaves are
reduced in size or may even fall to check transpiration. Thick cuticle on presence of wax
coating on exposed parts reduces cuticles transpiration.
Antitranspirants
A number of substances are known which when applied to the plants retard their
transpiration.

Such substances are called as antitranspirants. Some examples of

antitranspirants are colourless plastics, silicone, oils, low viscosity waxes, phenyl
mercuric acetate, abscisic acid, CO2, etc. Colourless plastic, silicone oils and low
38

viscosity waxes belong to one group as these are sprayed on the leaves, form after film
which is permeable to O2 and CO2 but not to water.
Fungicide phenyl mercuric acetate, when applied in low concentration (10 -4 m), it
exercised a very little toxic effect on leaves and resulted in partial closure of stomatal
pores for a period of two weeks. Similarly ABA a plant hormone also induces stomatal
closure. CO2 is an effective antitranspirants. A little rise in CO2 concentration from the
natural 0.03% to 0.05% induces partial closure of stomata. Its higher concentration
cannot be used which results in complete closure of stomata affecting adversely the
photosynthesis and respiration.

GUTTATION
In some plants such as garden nasturtium, tomato, colocasia etc, water drops ooze
out from the uninjured margins of the leaves where a main vein ends. This is called as
guttation and takes place usually early in the morning when the rate of absorption and
root pressure are high while the transpiration is very low.
The phenomenon of guttation is associated with the presence of special types of
stomata at the margins of the leaves which are called as water stomata or hydathodes.
Each hydathode consists of a water pore which remains permanently open.
Below this there is a small cavity followed by a loose tissue called as epithem.
This epithem is in close association with the ends of the vascular elements of veins.
Under high root pressure the water is given to the epithem by the xylem of the veins.
From epithem water is released into the cavity. When this cavity is completely filled with
watery solution, the later begins to ooze out in the form of watery drops through the
water pore.

39

Difference between transpiration and Guttation

Transpiration

Guttation

1. Water is lost from aerial parts of plants Watery solution oozes out from uninjured
in the form of invisible water vapours

margins of aerial leaves only

2. Transpiration occurs mostly through It occurs only through hydathodes (water

stomata. It may also takes place through stomata)
cuticle and lenticels
3. It takes place throughout the day, its rate It takes place only early in the morning
being maximum at noon.

when root pressure and the rate of water

absorption are higher

40

Lecture No: 7
MINERAL NUTRITION - INTRODUCTION - CRITERIA OF
ESSENTIALITY OF ELEMENTS - MACRO, SECONDARY AND
MICRONUTRIENTS - SOIL LESS CULTURE - SAND AND
HYDROPONICS
The term, mineral nutrient is generally used to refer to an inorganic ion obtained
from the soil and required for plant growth. The chemical form in which elements are
applied to plants is called as nutrient. Nutrition may be defined as the supply and
absorption of chemical compounds needed for plant growth and metabolism
The nutrients indispensable for the growth and development of higher plants are
obtained from three sources viz., atmosphere, water and soil. The atmosphere provides
carbon and oxygen as carbon dioxide. Carbon is reduced during photosynthesis and
oxygen is utilized during aerobic respiration. Soil provides the mineral ions.
Essential elements
The term essential mineral element was proposed by Arnon and Stout (1939).
These are the composition of both macro and microelements, in the absence of any one of
these elements the plant cannot maintain its normal growth and develops deficiency
symptoms, affects metabolism and die prematurely. Of the many elements that have been
detected in plant tissues, only 16 are essential for all higher plants. They are C, H, O, N,
P, K, Ca, Mg, S, Zn, Cu, Fe, Mn, B, Cl and Mo. In the absence of each of the essential
elements, plants develop deficiency symptoms characteristic of the deficient element and
die prematurely.
Macronutrients
The nutrient elements which are required for the growth of plants relatively in
larger quantities are called as major nutrients or macronutrients. The major elements
required for growth of plants are C, H, O, N, P, K, Ca, Mg and S. Among these nutrients,
C, H and O are taken up by the plants from the atmosphere and water. The N, P, K, Ca,
Mg and S are taken up by the plants from the soil and they are applied in the form of
chemical fertilizers either through the soil or foliage.
41

Micronutrients
The nutrient elements which are required comparatively in small quantities are
called as minor or micro nutrients or trace elements. The micronutrients required for the
plant growth are Zn, Cu, Fe, Mn, Mo, B and Cl.
Tracer elements or labeled elements
The nutrient elements that are required for plants are some times labeled and used
to study their movement or tracing out the involvement of such nutrients in metabolism in
different organs of plants, are called as tracer elements. They may either be stable or
radio active types and they are also called as isotopic elements.
E.g. Stable isotopes:
Radio active

15

N, 12C, 31P

: 14C, 32P, 65Zn, 56Fe, 60Co, etc.

Hidden hunger
When the plants are not able to meet their requirement either one or more of these
essential elements, the plants will undergo starvation for such elements. At the initial
stage of deficiency of such elements plants will not show any characteristic symptoms
which could be exhibited morphologically and due to want of those elements some
activities of plants would rather be affected and the internal deficiency is called as
Hidden hunger.
General role of essential elements
In general, an element is essential to the life of a higher green plant for one or
more of the following three reasons.
1. It may perform a nutritive role by being a component of one or more of the major
classes of plant constituents.
2. It may be a catalytic role either as an action for of an enzyme or as an integral
component of an enzyme.
3. It may function as a free ion and thereby exert a balancing role in maintaining
electro-neutrality within plant cells (e.g. Potassium).
Criteria for essentiality of elements
The demonstration of the essentially several elements (macro and micronutrients),
especially, micronutrients is rather very difficult. In view of the technical difficulties
associated with demonstrating the essentiality of elements required in very small

42

amounts, Arnon and Stout (1939) suggested the adoption of the following three criteria of
essentiality for judging the exact status of a mineral in the nutrient of a plant.
1. The element must be essential for normal growth or reproduction and the plant
processes cannot proceed without it.
2. The element cannot be replaced by another element.
3. The requirement must be direct i.e., not the result of some indirect effect such as
relieving toxicity caused by some other substance.
Another recent suggestion to the criteria of essentiality is that some elements
might better be called functional or metabolic elements rather than essential elements.
This is intended to indicate that an element that is metabolically active, functional or
metabolic may or may not be essential. For example in chlorine-bromine, chlorine is
designated as a functional element rather than an essential element as chlorine can be
substituted with bromine.
Based on the mobility in phloem, elements are also classified into three types.
1. Mobile elements

: N, K, P, S and Mg

2. Immobile elements

: Ca, Fe and B

3. Intermediate

: Zn, Mn, Cu, Mo

Functions of elements
Protoplasmic elements : N, P, S
Balancing elements

: Ca, Mg, K counteract to toxic effects of other minerals

by causing ionic balance.

Frame work elements

: C, H2O as they are the constituents of carbohydrates that

form cell walls.

Catalytic elements

: Mn, Cu, Mg, etc.

43

SOIL LESS GROWTH OR HYDROPONICS

The practice of growing plants in nutrient enriched water without soil is called as
soil less growth or hydroponics.

However, the term hydroponics is now being

applied to plants rooted in sand, gravel or other similar matter which is soaked with a
recycling flow of nutrient enriched water.
According to a recent limited nations report on hydroponics: In area of tropics,
where the water deficiency is the limiting factor in crop production, the soil less methods
hold out much promise because of the more economical use of water.
The report also indicated that in some areas, lack of fertile soil or very thin soil
layers may also move soil less methods worth serious consideration.
Besides these the other advantages of growing cucumbers, egg plants, peppers,
lettuces, spinach and other vegetables hydroponically under controlled environment are
1. The regulation of nutrients
2. Control of pets and diseases
3. Reduction of labour cost
4. Sometimes quicker yield
But there is two main drawbacks of hydroponics farming.
1. Firstly the cost of settling up the system is very high
2. Secondly it requires skills and knowledge its operation

44

Lecture No: 8
MECHANISM OF UPTAKE - PHYSIOLOGICAL ROLE OF NUTRIENTS
Mechanism
Previously it was thought that absorption of mineral salts takes place along with
water absorption.

But it is now understood that mineral salt absorption and water

absorption are two different processes.

Mineral salts are absorbed from the soil solution in the form of ions. They are
chiefly absorbed through the meristematic regions of the roots near the tips.
Plasma membrane of the root cells is not permeable to all the ions.

It is

selectivity permeable. All the ions of the same salt are not absorbed at equal rate but
leads unequal absorption of ions. First step in the absorption of mineral salts is the
process of Ion exchange which does not require metabolic energy.
The further processes of the absorption of mineral salts may be of two types.
1. Passive and 2. Active

1. Passive absorption
When the concentration of mineral salts is higher in the outer solution than in the
cell sap of the root cells, the mineral salts are absorbed according to the concentration
gradient by simple process of diffusion. This is called as passive absorption because it
does not require expenditure of metabolic energy.
Ion exchange
The ions adsorbed on the surface of the plasma membrane of the root cells may be
exchanged with the ions of same sign from external solution for eg. The cation K + of the
external soil solution may exchanged with H+ ions adsorbed on the surface of the plasma

45

membrane. Similarly anion may be exchanged with OH ions. There are two theories
regarding the mechanism of ion exchange.

1. Contact exchange theory

According to this theory the ions adsorbed or the surface of root cells and clay
particles are not held tightly but oscillate within small volume of space. If the roots and
clay particles are in close contact with each other, the oscillation volume of ions adsorbed
on root surface may over by the oscillation volume of ions adsorbed on clay particles, and
the ions adsorbed on clay particle may be exchanged with the ions adsorbed on root
surface directly without first being dissolved in soil solution.

2. Carbonic acid exchange theory

According to this theory, the CO 2 released during respiration of root cells
combines with water to form carbonic acid (H 2CO3). Carbonic acid dissociates into H+
and an anion HCO3 in soil solution. These H + ions may be exchanged for cations
adsorbed on the clay particles. The cations thus released into the soil solution from the
clay particles, may be adsorbed on root cells in exchange for H + ions or as in ion pairs
with bicarbonate.

Thus, the soil solution plays an important role in carbonic acid

exchange theory.

2. Active absorption of mineral salts

It has been observed that the cell sap in plants accumulates large quantities of
mineral slats ions against the concentration gradient. The accumulation of mineral salts
against to concentration gradient is an active process which involves the expenditure of
metabolic energy through respiration. The active absorption of mineral salts involves the
operation of a carrier compound present in the plasma membrane of the cells.

46

The carrier concept

According to this theory, the plasma membrane is impermeable to free ions. But
some compounds present in it acts as carrier and combines with ions to form carrier- ioncomplex which can move across the membrane. On the inner side of the membrane this
complex leaves releasing ions into the cell while the carrier goes back to the outer surface
to pick up fresh ions. They are two hypotheses based on the carrier concept to explain
the mechanism of active salt absorption. Although they are not universally accepted.
1. Lundegardhs cytochrome pump theory
Lundegardh and Burstrom (1933) believed that there was a definite correlation
between respiration and anion absorption. Thus when a plant is transferred from water to
a salt solution the rate of respiration increases. This increase in rate of respiration over
the normal respiration has been called as anion respiration or salt respiration.
Lundegardh (1954) proposed cytochrome pump theory which is based on the
following assumptions.
1. The mechanism of anion and cation absorption is different
2. Anions are absorbed through cytochrome chain by an active process.
(Cytochromes are ion porphyrin proteins that act as enzymes and helps in election
transfer during respiration).
3. Cations are absorbed passively.
According to this theory
1) Dehydrogenase reactions on inner side of the membrane give rise to protons (H +)
and electrons (e-).
2) The electrons travels over the cytochrome chain towards outside the membrane,
so that the Fe of the cytochrome becomes reduced (Fe++) on the outer surface and
oxidized (Fe+++) on the inner surface.
3) On the outer surface, the reduced cytochrome is oxidized by oxygen releasing the
electron (e-) and taking an anion (A-).
47

4) The electron thus released unites with H+ and oxygen to form water
5) The anion (A-) travels over the cytochrome chain towards inside.
6) On the inner surface the oxidized cytochrome becomes reduced by taking an
electron produced through the dehydrogenase reactions and the anion (A) is
released.
7) As the result of anion absorption, a cation (M) moves passively from outside to
inside to balance the anion.
2. Bennert Clarks protein Lecithin Theory
In 1856, Bennet Clark suggested that because the cell membranes chiefly
consist of phospholipids and proteins and certain enzymes seem to be located on them,
the carrier could be a protein associated with the phosphatide called as lecithin. He also
assumed the presence of different phosphatides to correspond with the number of known
competitive groups of cations and anions.
According to this theory
1. Phosphate group in the phosphatide is regarded as the active centre binding the
cations and the basic choline group as the anion binding centre.
2. The ions are liberated on the inner surface of the membrane by decomposition of
lecithin by the enzyme lecithinase.
3. The regeneration of the carrier lecithin form phosphatidic acid and choline takes
place in the presence of the enzyme choline acetylase and choline esterase and
ATP. The latter acts as a source of energy.
Donnans Equilibrium
The accumulation of ions inside the cells without involving expenditure of the
metabolic energy can be explained to some extent by Donnans equilibrium theory.
According to this theory there are certain pre existing ions inside the cell which
cannot diffuse outside through membrane. Such ions are called as in diffusible or fixed

48

ions. However, the membrane is permeable to both anions and cations of the outer
solutions.
Suppose there are certain fixed anions in the cell which is in contact with outer
solution containing anions and cations. Normally equal number of anions and cations
would have diffused into the cell through an electrical potential to balance each other, but
to balance the fixed anions more cations will diffuse into the cell. This equilibrium is
known as Donnans equilibrium. In this particular case, there would be an accumulation
of cations inside the cell.
If however, there are fixed cations inside the cell, the Donnans equilibrium will
result in the accumulation of anions inside the cell.

Specific roles of essential mineral elements

A. Macronutrients
1. Nitrogen

Nitrogen is important constituent of proteins, nucleic acids, porphyries

(chlorophylls & cytochromes) alkaloids, some vitamins, coenzymes etc

Thus N plays very important role in metabolism, growth, reproduction and

heredity.

2. Phosphorus

It is important constituent of nucleic acids, phospholipids, coenzymes NADP,

NADP H2 and ATP

Phospholipids along with proteins may be important constituents of cell

membranes

P plays important role in protein synthesis through nucleic acids and ATP

Through coenzymes NAD, NADP and ATP, it plays important role in energy
transfer reactions of cell metabolism eg. Photosynthesis, respiration and fat
metabolism etc.

49

Potassium

Although potassium is not a constituent of important organic compound in the

cell, it is essential for the process of respiration and photosynthesis

It acts as an activator of many enzymes involved in carbohydrate metabolism and

protein synthesis

It regulates stomatal movement

Regulates water balance

Calcium

It is important constituent of cell wall

It is essential in the formation of cell membranes

It helps to stabilize the structure of chromosome

It may be an activation of may enzymes

Magnesium

It is very important constituent of chlorophylls

It acts as activation of many enzymes in nucleic acid synthesis and carbohydrate

metabolism

It plays important role in binding ribosomal particles during protein synthesis.

Sulphur

It is important constituent of some amino acids (cystine, cysteine and methionine)

with which other amino acids form the protein

S helps to stabilize the protein structure

It is also important constituent of vitamin i.e biotin, thiamine and coenzyme A

Sulpho hydryl groups are necessary for the activity of many enzymes.

Iron

Important constituent of iron porphyrin proteins like cytochromes, peroxidase,

catalases, etc.

It is essential for chlorophyll synthesis

50

It is very important constituent of ferredoxin which plays important role in

photochemical reaction in photosynthesis and in biological nitrogen fixation.

Micro nutrients
Zinc

It is involved in the biosynthesis of growth hormone auxin (indole 3 acetic acid)

It acts activator of many enzymes like carbonic anhydrase and alcohol

dehydrogenase, etc.

Manganese

It is an activator of many respiratory enzymes

It is also an activator of the enzyme nitrite reductase

It is necessary for the evolution of oxygen (photolysis) during photosynthesis

Copper

It is an important constituent of plastocyanin (copper containing protein)

It is also a constituent of several oxidizing enzymes.

Boron

Boron facilitates the translocation of sugars by forming sugar borate complex.

It involves in cell differentiation and development since boron is essential for

DNA synthesis

Also involves in fertilization, hormone metabolism etc.

Molybdenum

It is constituent of the enzyme nitrate reductase and thus plays an important role
in nitrogen metabolism

It is essential for flower formation and fruit set.

51

Lecture No: 9

Foliar diagnosis - Nutritional and Physiological disorders foliar nutrition.

A. Foliar diagnosis - Symptoms
1. Nitrogen

Plant growth is stunted because protein content cell division and cell enlargement
are decreased

N deficiency causes chlorosis of the leave i.e yellowing older leaves are affected
first

In many plants eg. Tomato, the stem, petiole and the leaf veins become purple
coloured due to the formation of anthocyanin pigments.

2. Phosphorus

P deficiency may cause premature leaf fall

Dead necrotic areas are developed on leave or fruits

Leaves may turn to dark green to blue green colour. Sometimes turn to purplish
colour due to the synthesis and accumulation of anthocyanin pigments.

Potassium

Mottled chlorosis of leaves occurs

Neurotic areas develop at the tip and margins of the leaf

Plants growth remains stunted with shortening of internodes.

Calcium

Calcium deficiency causes disintegration of growing meristematic regions of root,

stem and leaves

Chlorosis occurs along the margins of the younger leaves

52

Malformation of young leaves takes place

Magnesium

Mg deficiency causes mottled chlorosis with veins green and leaf tissues yellow
or white appearing first on older leaves

Dead neurotic patches appear on the leaves

In cotton Mg deficiency leads o reddening of leaves and disorder is called as

reddening in cotton.

Sulphur

Deficiency causes chlorosis of the leaves

Tips and margins of the leaf roll in ward

Stem becomes hard due to the development of sclerenchyma.

Micronutrients
Iron
Iron deficiency causes chlorosis of young leaves which is usually interveinal.
Zinc

Zinc deficiency causes chlorosis of the young leaves which starts from tips and
the margins

The size of the young leaves is very much reduced. This disorder is called as
little leaf disease

Stalks will be very short.

Manganese

The young leaves are affected by mottled chlorosis

Veins remain green

Small necrotic spots developed on the leaves with yellow strips

Copper

Copper deficiency causes necrosis of the tip of the young leaves

It also causes die-back of citrus and fruit trees

53

Also causes reclamation disease or white tip disease of cereals and leguminous
plants.

Boron

Boron deficiency causes death of shoot tip

Flower formation is suppressed

Root growth is stunted

The other diseases caused by B deficiency is

Heart rot of beet

Stem crack of celery

Brown heart of cabbage

Water core of turnip

Internal cork formation in apple

Hen and chicken in grapes

Molybdenum

Molybdenum deficiency causes interveinal chlorosis of older leaves

Flower formation is inhibited

Causes whiptail disease in cauliflower plants.

Foliar Nutrition
Foliar nutrition is fertilizing certain crop plants through aerial spraying.
Mechanism
Penetration of the spray solution or nutrient solution occurs through cuticle the
layer of polymerized wax which occurs on outer surface of the epidermal cells of leaves.
After penetration in the cuticle, further penetration take place through fine, thread
like semi-microscopic structure called ectodesmata.

This extends through the outer

epidermal cell wall, from the inner surface of the cuticle to the plasma membrane.

54

When the substance reaches plasma membrane of an epidermal cell, it will be

observed by mechanism similar to those which operate in root cells.

Advances
1. Foliar nutrition may serve as a mean of applying supplemental macronutrients
during critical growth periods when it is impracticable to apply fertilizers to soil.
Eg. Unusual period of dry weather.
2. Foliar nutrition may afford a remedy for the time lag between soil applied and
plant absorbed. Time is too long because of fast growing rates.
NUTRITIONAL DISORDERS
When a nutrient element insufficiency (deficiency and/or toxicity) occurs, visual
symptoms may or may not appear, although normal plant development will be slowed.
When visual symptoms do occur, such symptoms can frequently be used to identify the
source of the insufficiency.
Deficiency Symptoms

Stunted or reduced growth of the entire plant with the plant itself either remaining
green or lacking an over-all green color with either the older or younger leaves
being light green to yellow in color.

Chlorosis of leaves, either interveinal or of the whole leaf itself, with symptoms
either on the younger and/or older leaves, or both (chlorosis due to the loss or lack
of chlorophyll production).

Necrosis or death of a portion (margins or interveinal areas) of a leaf, or the whole

leaf, usually occurring on the older leaves.

Slow or stunted growth of terminals (rosetting), the lack of terminal growth, or

death of the terminal portions of the plant.

A reddish purpling of leaves, frequently more intense on the under side of older
leaves due to the accumulation of anthocyanin (Mottling)

55

Chlorosis is caused by the deficiency of mineral elements such as Mn, K, Zn, Fe, Mg, S
and N. Mottling is caused due to the deficiencies of N, Mg, P, S and Necrosis due to the
deficiency of Mg, K, Zn, Ca and Mo.
Toxicity Symptoms
Visual symptoms of toxicity may not always be the direct effect of the element in
excess on the plant, but the effect of the excess element on one or more other elements.
For example, an excessive level of potassium (K) in the plant can result in either
magnesium (Mg) and/or calcium (Ca) deficiency, excess phosphorus (P) can result in a
zinc (Zn) deficiency and excess Zn in an iron (Fe) deficiency.
These effects would compare to elements, such as boron (B), chlorine (Cl), copper
(Cu), and manganese (Mn), which create visual symptoms that are the direct effect of an
excess of that element present in the plant.
Some elements, such as aluminum (Al) and copper (Cu) can affect plant growth
and development due to their toxic effect on root development and function.
Hidden Hunger
In some instances, a nutrient element insufficiency may be such that no symptoms
of stress will visually appear with the plant seeming to be developing normally. This
condition has been named hidden hunger, a condition that can be uncovered by means of
either a plant analysis and/or tissue test.
A hidden hunger occurrence frequently affects the final yield and the quality of
the product produced. For grain crops, the grain yield and quality may be less than
expected; for fruit crops, abnormalities, such as blossomed rot and internal abnormalities
may occur, and the post harvest characteristics of fruits and flowers will result in poor
shipping quality and reduced longevity. Another example is potassium (K) insufficiency
in corn, a - deficiency that is not evident until at maturity when plants easily
PHYSIOLOGICAL DISORDERS
Physiological disorder is the abnormal growth pattern or abnormal external or
internal conditions of fruits due to adverse environmental conditions such as deviation
from normal state of temperature, light, moisture, nutrient, harmful gases and inadequate
supply of growth regulators.
56

Disorders associated with low temperature

1. Leaf chlorosis and frost banding
Chlorosis was caused by a disruption of chloroplasts caused by winter cold. Green
chlorophyll pigments are often converted in to yellow pigment. Leaf may appear with
distinct bleached bands across the blade of young plants called frost banding e.g.:
sugarcane, wheat and barley.
2. Leaf necrosis and malformations
Spring frost causes various types and degree of injury including cupping, crinkling
finishing and curling of leaves of apple trees and stone fruits. The distortion is caused by
death of the developed tissues before the expansion of leaves.
3. Stem disorders
Frost cracks develop when tree trunk or limps lost their heat too rapidly. The outer
layer of bark and wood cool most rapidly and subjected to appreciable tension causing
marked shrinkage and cracking following a sudden temperature drop. Affected timber is
of poor quality.
Disorders associated with high temperature
1. Leaf scorch
High temperature causes leaf scorch directly or indirectly by stimulating excessive
evaporation and transpiration. Tip burn of potato is a widespread example for this
disorder.
2. Sunscald
In leaf vegetable crops like lettuce and cabbage, when leaves on the top of the head
are exposed to intense heat, water soaked lesions or blistered appearance occur These
irregular shaped areas become bleached and parched later.
3. Water core
In fruit crop like Tomato, exposure to high temperature causes death of the
outer cells of fruit skin. Subsequently corky tissue occurs beneath the skin, with watery
appearance of the flesh near the core of the fruits faster. Often light stress is coupled with

57

heat stress e.g. sun scald of bean, sun burning of soybean and cowpea. In flower crop like
chrysanthemum, increase in light intensity affects flower bud formation. Reproduction
phase does not commence and modified into leaf like bracts.
Disorders caused by light stress
Adverse light intensity causes impaired growth and reduced vigour. Subsequently
leaves gradually lose green colour, turning pale green to yellow, stems may dieback little
every year. Insufficient light limits photosynthesis, causing food reserves to be depleted.
Identification of Physiological Disorders and Corrective Measures
Crop
Rice

Malady

Corrective measure
1%super phosphate

Severe chlorosis of leaves

and

0.5%

ferrous sulphate
Rice

Irregular flowering and chaffiness 1% super phosphate and magnesium

Rice

multiple deficiency of nutrients

sulphate.
Tip drying and marginal scoring 1%super phosphate and 0.5% zinc

Maize

and browning
Chlorosis

Maize

ferrous sulphate and 0.5% urea.

'White bud' yellowing in the bud 0.5% zinc sulphate spray with

Maize

leaves only
1% urea.
Tip drying and marginal scoring 1% super phosphate and 0.5% zinc

Maize
Sorghum

sulphate.
A spray solution containing 0.5%

pinkish colouration of lower leaves

Marginal scorching and yellowing.
Irregular drying of tips and margins
Chlorosis of younger leaves

sulphate.
0.5% ferrous sulphate and 1%urea
25 kg of zinc sulphate / ha
Spray of 0.5% ferrous sulphate with
0.5%urea and 0.5% ammonium
sulphate

Pulses
Cowpea

Water soaked necrotic spots on leaf Spray containing sulphate and zinc
surface.
restricted

Root growth very much sulphate 0.1% and 0.1% urea

in

10-12

days

seedling
58

old

Oilseeds
Groundnut

Chlorosis of terminal leaves

0.5% ferrous sulphate and urea 1%

59

Lecture No: 10
PHOTOSYNTHESIS - REQUIREMENTS OF PHOTOSYNTHESIS LIGHT, CO2, PIGMENTS AND H20
Photosynthesis is a vital physiological process where in the chloroplast of green
plants synthesizes sugars by using water and carbon dioxide in the presence of light.
Photosynthesis literally means synthesis with the help of light i.e. plant synthesize organic
matter (carbohydrates) in the presence of light.
Photosynthesis is sometimes called as carbon assimilation (assimilation:
absorption into the system).This is represented by the following traditional equation.
Light
6CO2 + 12H2O

6O2 + C6H12O6
Green pigments

carbohydrates

During the process of photosynthesis, the light energy is converted into chemical
energy and is stored in the organic matter, which is usually the carbohydrate. One
molecule of glucose for instance, contains about 686 K Calories energy. CO 2 and water
constitute the raw material for this process and oxygen and water are formed as the by
products during photosynthesis. Stephen Hales (1727) first explained the relationship
between sunlight and leaves and Sachs (1887) established that starch was the visible
product of photosynthesis.
Photosynthetic apparatus
The chloroplast in green plants constitutes the photosynthetic apparatus. In higher
plants, the chloroplast is discoid in shape, 4-6 in length and 1-2 thick. The chloroplast
is bounded by two unit membranes of approximately 50A thickness and consists of
lipids and proteins. The thickness of the two membranes including the space enclosed by
them is approximately 300A (1 Angstrom: 0.1 cm).
Internally, the chloroplast is filled with a hydrophilic matrix called as stroma
embedded with grana.

Each grana consists of 5-25 disk shaped grana lamellae

(thylakoid) placed one above the other like the stack of coins. Each grana lamella of
thylakoid encloses a space called loculus and the thylakoid membrane consists of
60

alternating layer of lipids and proteins. Some of the grana lamella of thylakoid of grana
are connected with thylakoid of other grana by somewhat thinner stroma lamella or fret
membrane. Chlorophyll and other photosynthetic pigments are confined to grana. The
chlorophylls are the site of photochemical reactions.
Photosynthetic pigments
Photosynthetic pigments are of three types; Chlorophylls, Carotenoids and Phycobillins.

Chlorophylls and Carotenoids are insoluble in water and can be extracted only
with organic solvents such as acetone, petroleum ether and alcohol.

Phycobillins are soluble in water

Carotenoids include carotenes and xanthophylls. The xanthophylls are also called
as carotenols.

Chlorophylls (green pigments)

Chlorophylls are magnesium porphyrin compounds. The porphyrin ring consists
of four pyrrol rings joined together by CH bridges. Long chain C atoms called as phytol
chain is attached to porphyrin ring at pyrrol ring IV.
The chemical structure of chlorophyll a and chlorophyll b are well established. The
molecular formula for chlorophyll a: C55H72O5N4 Mg and chlorophyll b: C55H70O6N4 Mg.
Both of them consist of Mg porphyrin head which is hydrophilic and a phytol tail which
is lipophilic. The two chlorophylls differ because in chlorophyll b there is a CHO group
instead of CH3 group at the 3rd C atom in pyrrol ring II.
Chlorophyll is formed from protochlorophyll in light. The protochlorophyll lacks
2H atoms one each at 7th and 8th C atoms in pyrrol ring IV.
Carotenoids (yellow or orange pigments)
1. Carotenes: Carotenes are hydrocarbons with a molecular formula C40H56
2. Xanthophylls (carotenols)
They are similar to carotenes but differ in having two oxygen atoms in the form of
hydroxyl or carboxyl group. The molecular formula is C 40H56O2.The role of Carotenoids
is absorption of light energy and transfer the light energy to chlorophyll a molecules.
61

They also play a very important role in preventing photodynamic damage within the
photosynthetic apparatus. Photodynamic damage is caused by O2 molecules which is very
reactive and is capable of oxidizing whole range of organic compounds such as
chlorophylls and there by making them unfit for their normal physiological function.
Phycobillins (red and blue pigments)
These also contain four pyrrol rings but lack Mg and the phytol chain.
Location of photosynthetic pigments in chloroplast
The photosynthetic pigments are located in grana portions of the chloroplast.
They are present in the thylakoid membrane or membrane of grana lamella. The
membrane of thylakoid is made up of proteins and lipids or the membrane consists of
both lipid layer and protein layer. The hydrophilic heads of the chlorophyll molecules
remain embedded in the protein layer while lipophilic phytol tail in the lipid layer. The
other pigments are thought to be present along with chlorophyll molecules.
Distribution of photosynthetic pigments in plant kingdom
Pigments
Chlorophylls
Chlorophyll a
Chlorophyll b
Chlorophyll c
Chlorophyll d
Bacteria chlorophylls
a, b, c, d & e
Carotenoids
Carotenes ( and )
Xanthophylls
Lutein
Violaxanthin
Fucoxanthin
Phycobillins
Phycocyanins
Phycoerythrins
Allophycocyanin

Distribution in plant kingdom

All photosynthesizing plants except bacteria
Higher plants and green algae
Diatoms and brown algae
Red algae
Purple and green bacteria

Higher plants and algae

Higher plants and algae
Green leaves and Green and Red algae
Green leaves
Brown algae
Blue green algae and red algae
Blue green algae and Red algae
Blue green and Red algae

62

Light
The chief source of light energy for photosynthesis is sun. The solar radiation or
solar energy passes through the space and reaches the earth in the form of
electromagnetic radiation with waves of varying lengths.

The various portions of

electromagnetic spectrum are gamma rays, ultraviolet rays, visible rays and infrared rays.
The wavelength of these rays ranges from 280 nm to 1000 nm.

Below 280 nm
- X rays, Gamma rays and Cosmic rays
280-390 nm
- Ultra violet radiation
400-510 nm
- Blue light
510-610 nm
- Green light
Visible light (PAR)
610-700 nm
- Red light
(VIBGYOR)
700-1000 nm
- Far red light (IR)
Photosynthetic pigments absorb light energy only in the visible part of the
spectrum. The earth receives only about 40% (or about 5x10 20 K cal) of the total solar
energy. The rest is either absorbed by the atmosphere or scattered into the space. Only
about 1% of the total solar energy received by the earth is absorbed by the pigments and
utilized in photosynthesis.
Absorption spectra of chlorophyll
The absorption of different wavelengths of light by a particular pigment is called
absorption spectrum. Chlorophylls absorb maximum light in the violet blue and red part

63

of the spectrum. The absorption peaks of chlorophyll a are 410 and 660; for chlorophyll
b 452 and 642. Carotenoids absorb light energy in blue and blue green part of the
spectrum.

64

65

Transfer of light energy absorbed by accessory pigments to chlorophyll a

All pigments except chlorophyll a are called as accessory pigments or antenna
pigments. The light energy absorbed by accessory pigments is transferred to chlorophyll
a molecule. The transfer of light energy from accessory pigments to chlorophyll a is
called as resonance or Forster transfer and takes part in primary photochemical reaction
in photosynthesis. Chlorophyll a molecules also absorb light energy directly. As a result
of absorbing the light energy, the chlorophyll molecule gets excited.
Excited states of atoms or molecules (fluorescence and phosphorescence)
The normal state of the chlorophyll molecule or atom is called as ground state or
singlet state. When an electron of a molecule or an atom absorbs a quantum of light, it is
raised to a higher energy level which is called as excited second singlet state. This state
is unstable and has a life time of 10-12 seconds.
The electron comes to the next higher energy level by the loss of some of its extra
energy in the form of heat. This higher energy level is called as excited first singlet state
and is also unstable with a half life of 10-9 seconds. From the first singlet state, the excited
electron may return to the ground state in two ways viz., either losing its remaining extra
energy in the form of heat or in the form of radiant energy. The second process is called
fluorescence. The chlorophyll molecules exit the extra energy in the form of fluorescent
light when they are exposed to incident light. Fluorescent light is of longer wavelength
than the incident light.
The excited molecule or the atom may also lose its excitation energy by internal
conversion and comes to another excited state called as triplet state which is meta stable
with a half life of 10-3 seconds. From the triplet state, the excited molecule or the atom
may return to the ground state in three ways.
(i)

By losing its remaining extra energy in the form of heat

(ii)

By losing extra energy in the form of radiant energy (phosphorescence) and

the chlorophyll molecules emit phosphorescent light even after the incident
radiant light is cut off. The phosphorescent light is of longer wavelength than
incident light and also fluorescent light.

66

(iii)

Electrons carrying the extra energy may be expelled from the molecule and is
consumed in some further photochemical reaction and the fresh normal
electron returns to the molecule.

Quantum requirement and quantum yield

Light rays consist of tiny particles called photons and the energy carried by a
photon is called quantum. The number of photons (quantum) required to release one
molecule of oxygen in photosynthesis is called quantum requirement. On the other hand,
the number of oxygen molecules released per photon of light in photosynthesis is called
as quantum yield. The quantum yield is always in fraction of one.
Warburg found minimum quantum requirement for photosynthesis as four. It is
because the reduction of one molecule of CO2 by two molecules of H2O requires the
transfer of 4H atoms. The transfer of each H atoms from H2O to CO2 requires one photon
or quantum of light.
4H2O

4 OH- + 4H+

4OH-

2H2O + O2 + 4e-

4H+ + CO2

(CH2O) + H2O

2H2O + CO2

(CH2O) + O2 + H2O

(CH2O) in the above equation represent 1/6 of the carbohydrate molecule such as
glucose. One molecule of glucose contains 686 K. cal of energy. Therefore, 1/6 glucose
molecule contains 686/6 i.e., approximately 112 K.cal energy. It is also known that the
rate of photosynthesis is maximum at red light and each photon of red light contains
about 40 K cal. of energy. This would suggest that the efficiency with which the plants
can convert light energy into chemical energy is 112 / 40 x 4: 70%, which indeed is very
high.
According to Emerson and his coworkers, photosynthesis is a very complicated
process and is not so efficient to convert all the light energy into chemical energy. There
is a considerable loss of light energy absorbed during photosynthesis and therefore the
minimum quantum requirement for photosynthesis as suggested by Emerson and
coworkers are 8-10. Considering that the quantum requirement for photosynthesis is 810, the quantum yield would accordingly be 1/8 to 1/10 (0.125 to 0.10)
67

Lecture No: 11
MECHANISM OF PHOTOSYNTHESIS - LIGHT REACTION - CYCLIC
AND NON CYCLIC PHOTOPHOSPHORILATION - RED DROP EMERSON ENHANCEMENT EFFECT

Photo systems (Two pigment systems)

The discovery of red drop and the Emersons enhancement effect led the scientists
to suggest that photosynthesis is driven by two photochemical processes. These processes
are associated with two groups of photosynthetic pigments called as pigment system I and
pigment system II. Wavelength of light shorter than 680 nm affect both the pigments
systems while wavelength longer than 680 nm affect only pigment system I.
In green plants, pigment system I contains chlorophyll a, b and carotene. In this
pigment system, a very small amount of chlorophyll a absorbing light at 700 nm, known
as P700 however constitutes the reaction centre of photosystem I.
The pigment system II contains chlorophyll b and some forms of chlorophyll a
(such as chlorophyll a 662, chlorophyll a 677 and chlorophyll a 679) and xanthophylls.
A very small amount of special form of chlorophyll called P680 constitute the reaction
centre of pigment system II. Carotenoids are present in both the pigment systems
The two pigment systems I and II are interconnected by a protein complex called
cytochrome b6f complex. The other intermediate components of electron transport chain
viz., plastoquinone (PQ) and plastocyanin (PC) act as mobile electron carriers between
the complex and either of the two pigment systems. The light energy absorbed by other
pigment is ultimately trapped by P700 and P680 forms of chlorophyll a which alone take
part in further photochemical reaction.
Pigment system I (PSI) complex consists of 200 chlorophylls, 50 Carotenoids and
a molecule of chlorophyll a absorbing light at 700 nm(P700) and this constitute the
reaction centre of photosystem I. Pigment system II (PSII) complex consists of 200
chlorophylls, 50 Carotenoids and a mole of chlorophyll a absorbing light at 680 nm,
called P 680 at the centre. This constitutes the reaction centre of pigment system II.

68

Photosynthetic units the Quantasomes

Emerson and Arnold (1932) showed that about 2500 chlorophyll molecules are
require fixing one molecule of CO2 in photosynthesis. This number of chlorophyll
molecules was called the chlorophyll unit but the name was subsequently changed to
photosynthetic unit. However, since the reduction or fixation of one CO 2 molecule
requires about 10 quanta of light, it is assured that 10 flashes of light are required to yield
one O2 molecule or reduction of one molecule of CO 2. Thus each individual unit would
contain 1/10th of 2500 i.e., 250 molecules.
Action spectrum
The pigments present in plants or any living organism have the ability to absorb
radiant energy to carry out photo physiological reactions. It is difficult to decide which
specific pigment is actually associated with the particular photochemical reactions.
Hence, a common procedure to identify the pigment involved in a particular
photoreaction is to determine the action spectrum i.e. measuring the rate of the particular
photoreaction. Once the action spectrum for a photo physiological reaction is determined,
the next step is to compare this action spectrum with absorption spectrum of a pigment.
Two pigments, A and B were isolated from the same plant and their absorption
spectra were determined. Pigment A has a peak in absorption at 395 nm and the pigment
B at 660 nm. The close correspondence between the absorption spectrum and the action
spectrum of pigment B strongly supports that Pigment B is responsible for absorbing
radiant energy to drive this photoreaction.
Mechanism of photosynthesis
The biosynthesis of glucose by the chloroplast of green plants using water and
CO2 in the presence of light is called photosynthesis. Photosynthesis is a complex process
of synthesis of organic food materials. It is a complicated oxidation- reduction process
where water is oxidized and CO2 is reduced to carbohydrates. The mechanism of
photosynthesis consists of two parts.
1. Light reaction / Primary photochemical reaction / Hills reaction/ Arnons cycle
2. Dark reaction / Black mans reaction / Path of carbon in photosynthesis.

69

1. Light reaction or Primary photochemical reaction or Hills reaction

In light reaction, ATP and NADPH2 are produced and in the dark reaction, CO2 is
reduced with the help of ATP and NADPH 2 to produce glucose. The light reaction is
called primary photochemical reaction as it is induced by light. Light reaction is also
called as Hills reaction as Hill proved that chloroplast produce O2 from water in the
presence of light. It is also called as Arnons cycle because Arnon showed that the H+ ions
released by the break down of water are used to reduce the coenzyme NADP to NADPH.
Light reaction includes photophosphorylation as ATP is synthesized in the presence of
light. The reaction takes place only in the presence of light in grana portion of the
chloroplast and it is faster than dark reaction. The chlorophyll absorbs the light energy
and hence the chlorophyll is called as photosystem or pigment system. Chlorophylls are of
different types and they absorb different wavelengths of light. Accordingly, chlorophylls
exist in two photo systems, Photosystem I (PSI) and Photosystem II (PS II). Both photo
systems are affected by light with wavelengths shorter than 680nm, while PS I is affected
by light with wavelengths longer than 680nm.
The components of photo systems
Photosystem I
Chlorophyll a 670

Photosystem II
Chlorophyll a 660

Chlorophyll a 680

Chlorophyll a 670

Chlorophyll a 695

Chlorophyll a 680 or P680

Chlorophyll a 700 or P700

Chlorophyll b

Chlorophyll b

Phycobillins

Carotenoids

Xanthophylls

P700 form of Chlorophyll a

P680 form of Chlorophyll a

is the active reaction centre

is the active reaction centre

The light reaction can be studied under the following headings.

i. Absorption of light energy by chloroplast pigments
Different chloroplast pigments absorb light in different regions of the visible part
of the spectrum.
ii. Transfer of light energy from accessory pigments to chlorophyll a
70

All the photosynthetic pigments except chlorophyll a are called as accessory or

antenna pigments. The light energy absorbed by the accessory pigments is transferred by
resonance to chlorophyll a which alone can take part in photochemical reaction.
Chlorophyll a molecule can also absorb the light energy directly. In pigment system I, the
photoreaction centre is P700 and in pigment system II, it is P680.
iii. Activation of chlorophyll molecule by photon of light
When P700 or P680 forms of chlorophyll a receives a photon (quantum) of light,
becomes an excited molecule having more energy than the ground state energy. After
passing through the unstable second singlet state and first singlet stage the chlorophyll
molecules comes to the meta stable triplet state.

This excited state of chlorophyll

molecule takes part further in primary photochemical reaction i.e. the electron is expelled
from the chlorophyll a molecule.
Light
Chlorophyll a

Excited triplet state of chlorophyll a

Chlorophyll a+ + e-

Excited triplet state of chlorophyll a

iv. Photolysis of water and O2 evolution (oxidation of water)

These processes are associated with pigment system II and are catalyzed by Mn ++
and Cl- ions. When pigment system II is active i.e it receives the light, the water
molecules split into OH- and H+ ions (Photolysis of water). The OH- ions unite to form
some water molecules again and release O2 and electrons.
4H2O

4H+ + 4 (OH-)

4(OH-)

2H2O + O2 + 4e-

2H2O

4H+ + O2 + 4e-

v. Electron transport and production of assimilatory powers (NADPH2 and ATP)

It has already been observed that when chlorophyll molecule receives the photon
of light, an electron is expelled from the chlorophyll a molecule along with extra energy.
This electron after traveling through a number of electron carriers is utilized for the
production of NADPH2 from NADP and also utilized for the formation of ATP molecules
from ADP and inorganic phosphate (Pi). The transfer of electrons through a series of
71

coenzymes is called electron transport and the process of formation of ATP from ADP
and Pi using the energy of electron transport is called as photosynthetic phosphorylation
or photophosphorylation. The types of Phosphorylation include cyclic and non- cyclic.

Cyclic electron transport and cyclic photophosphorylation

The electrons released from photosystem I goes through a series of coenzymes

and returns back to the same photosystem I. This electron transport is called cyclic
electron transport. The synthesis of ATP occurring in cyclic electron transport is called
cyclic photophosphorylation. The cyclic electron transport involves only pigment system
I. This situation is created when the activity of pigment system II is blocked. Under this
condition,
1. Only pigment system I remain active
2. Photolysis of water does not take place
3. Blockage of noncyclic ATP formation and this causes a drop in CO2 assimilation in

72

dark reaction
4. There is a consequent shortage of oxidized NADP
Thus, when P700 molecule is excited in pigment system I by absorbing a photon
(quantum) of light, the ejected electron is captured by ferredoxin via FRS.

From

ferredoxin, the electrons are not used up for reducing NADP to NADPH + H + but
ultimately it falls back to the P700 molecule via number of other intermediate electron
carriers. The electron carriers are probably cytochrome b6, cytochrome f and
plastocyanin.
During this electron transport, phosphorylation of ADP molecule to form ATP
molecule take place at two places i.e., between ferredoxin and cytochrome b6 and
between cytochrome b6 and cytochrome f. Thus, two ATP molecules are produced in this
cycle. Since the electron ejected from P700 molecule is cycled back, the process has been
called as cyclic electron transport and the accompanying phosphorylation as the cyclic
photophosphorylation.

73

Significance of cyclic photophosphorylation

1. During cyclic electron transport and phosphorylation, photolysis of water, O 2
evolution and reduction of NADP do not take place.
2. The electron returns or cycles back to original position in the P700 form of
chlorophyll a. Here, chlorophyll molecule serves both as donor and acceptor of
the electron.
3. It generates energy rich ATP molecules at two sites and as such cannot drive dark
reactions of photosynthesis
On the other hand, non- cyclic photophosphorylation does not produce sufficient
ATP in relation to NADPH to operate the dark phase of photosynthesis. Therefore, the
deficiency of ATP molecule in noncyclic photophosphorylation is made up by the
operations of cyclic photophosphorylation.
Secondly, the cyclic photophosphorylation may be an important process in
providing ATP for photosynthesis and other processes such as synthesis of starch,
proteins, lipids, nucleic acids and pigments within the chloroplast.
Non cyclic photophosphorylation
The electron released from photosystem II goes through a series of enzymes and
Co-enzymes to photosystem I. This is called non cyclic electron transport and the
Synthesis of ATP in non cyclic electron transport is called non- cyclic photo
phosphorylation. The main function of non cyclic electron transport is to produce the
assimilatory powers such as NADPH2 and ATP and the process occurs in photosystem I
and II.
This process of electron transport is initiated by the absorption of a photon
(quantum) of light by P680 form of chlorophyll a molecule in the pigment system II,
which gets excited and an electron is ejected from it so that an electron deficiency or a
hole is left behind in the P680 molecule.
The ejected electron is trapped by an unknown compound known as Q. From Q,
the electron passes downhill along a series of compounds or intermediated electron
carriers such as cytochrome b6, plastoquinone, cytochrome f and a copper containing

74

plastocyanin and ultimately received by pigment system I. At one place during electron
transport i.e. between plastoquinone and cytochrome f, one molecule of ATP is formed
from ADP and inorganic phosphate.
Now, when a photon of light is absorbed by P700 form of chlorophyll molecule in
the pigment system I, this gets excited and an electron is ejected from it. This ejected
electron is trapped by FRS (Ferredoxin Reducing Substance) and it is then transferred to
a non-heme iron protein called ferredoxin. From ferredoxin, electron is transferred to
NADP so that NADP is reduced to NADPH + H+
The hole in pigment system I has been filled by electron coming from pigment
system II. But, the hole or an electron deficiency in pigment system II is filled up by the
electron coming from photolysis of water where, water acts as electron donor.
In this scheme of electron transport, the electron ejected from pigment system II
did not return to its place of origin, instead it is taken up by pigment system I. Similarly,
the electron ejected from pigment system I did not cycle back and was consumed in
reducing NADP. Therefore, this electron transport has been called as noncyclic electron
transport and accompanying phosphorylation as noncyclic photophosphorylation.
The non cyclic electron transport (photophosphorylation) takes the shape of Z and
hence it is called by the name Zscheme. Non cyclic photophosphorylation and O 2
evolution are inhibited by CMU (3-(4-Chlorophyl) 1-1dimethyl urea and 3-(3-4dichlorophenyl)-1, 1-dimethyl urea (DCMU).
Significance of non cyclic electron transport
1.

It involves PS I and PSII

2.

The electron expelled from P680 of PSII is transferred to PS I and hence it is a

non cyclic electron transport.

3.

In non cyclic electron transport, photolysis of water (Hills reaction and evolution
of O2) takes place.

4.

Phosphorylation (synthesis of ATP molecules) takes place at only one place.

5.

The electron released during photolysis of water is transferred to PS II.

6.

The hydrogen ions (H+) released from water are accepted by NADP and it
becomes NADPH2

75

7.

At the end of non cyclic electron transport, energy rich ATP, assimilatory power
NADPH2 and oxygen from photolysis of water are observed.

8.

The ATP and NADPH2 are essential for the dark reaction wherein, reduction of
CO2 to carbohydrate takes place.

Comparison of cyclic and non cyclic electron transport and photophosphorylation in

chloroplasts

1
2

Cyclic electron transport and photo

Non cyclic electron transport and

phosphorylation in chloroplasts

photo phosphorylation in chloroplasts

Associated with pigment system I

The electron expelled from chlorophyll

Associated with pigment system I and II

The electron expelled from chlorophyll

molecule is cycled back

molecule is not cycled back. But, its loss is

compensated by electron coming from

photolysis of water
Photolysis of water and evolution of O2 Photolysis of water and evolution of O2

do not take place

Phosphorylation takes place at two

take place
Phosphorylation takes place at only one

places
NADP + is not reduced

place
NADP + is reduced to NADPH+ + H+

Significance of light reaction

1.

Light reaction takes place in chlorophyll in the presence of light.

2.

During light reaction, the assimilatory powers ATP and NADPH2 are synthesized.

3.

The assimilatory powers are used in dark reaction for the conversion of CO2 into
sugars.

4.

Photolysis of water occurs in light reaction. The H+ ions released from water are
used for the synthesis of NADPH2

5.

Plants release O2 during light reaction

Red drop and Emersons enhancement effect

76

Robert Emerson noticed a sharp decrease in quantum yield at wavelength greater

than 680 nm, while determining the quantum yield of photosynthesis in chlorella using
monochromatic light of different wavelengths. Since this decrease in quantum yield took
place in the red part of the spectrum, the phenomenon was called as red drop.
Later, they found that the inefficient far-red light beyond 680 nm could be made
fully efficient if supplemented with light of shorter wavelength (blue light). The quantum
yield from the two combined beams of light was found to be greater than the sum effects
of both beams used separately. This enhancement of photosynthesis is called as
Emersons Enhancement.

77

Lecture No: 12
PHOTOSYNTHETIC PATHWAYS - C3, C4 AND CAM
Dark reaction or Blackmans reaction or Path of carbon in photosynthesis
This is the second step in the mechanism of photosynthesis. The chemical
processes of photosynthesis occurring independent of light is called dark reaction. It
takes place in the stroma of chloroplast. The dark reaction is purely enzymatic and it is
slower than the light reaction. The dark reactions occur also in the presence of light. In
dark reaction, the sugars are synthesized from CO2. The energy poor CO2 is fixed to
energy rich carbohydrates using the energy rich compound, ATP and the assimilatory
power, NADPH2 of light reaction. The process is called carbon fixation or carbon
assimilation. Since Blackman demonstrated the existence of dark reaction, the reaction is
also called as Blackmans reaction. In dark reaction two types of cyclic reactions occur
1. Calvin cycle or C3 cycle
2. Hatch and Slack pathway or C4 cycle
Calvin cycle or C3 cycle
It is a cyclic reaction occurring in the dark phase of photosynthesis. In this
reaction, CO2 is converted into sugars and hence it is a process of carbon fixation. The
Calvin cycle was first observed by Melvin Calvin in chlorella, unicellular green algae.
Calvin was awarded Nobel Prize for this work in 1961. Since the first stable compound in
Calvin cycle is a 3 carbon compound (3 phosphoglyceric acid), the cycle is also called as
C3 cycle. The reactions of Calvins cycle occur in three phases.
1. Carboxylative phase
2. Reductive phase
3. Regenerative phase

78

1. Carboxylative phase
Three molecules of CO2 are accepted by 3 molecules of 5C compound viz.,
ribulose diphosphate to form three molecules of an unstable intermediate 6C compound.
This reaction is catalyzed by the enzyme, carboxy dismutase
3 CO2

3 Ribulose
diphosphate

Carboxy dismutase 3 unstable intermediate 6

carbon compound

The three molecules of the unstable 6 carbon compound are converted by the addition of
3 molecules of water into six molecules of 3 phosphoglyceric acid. This reaction is also
catalyzed by the enzyme carboxy mutase.
3 unstable
intermediate 6 C
compound

3 H2O

Carboxy
dismutase

3 phosphoglyceric acid

3 phosphoglyceric acid (PGA) is the first stable product of dark reaction of

photosynthesis and since it is a 3 carbon compound, this cycle is known as C3 cycle.

79

2. Reductive phase
Six molecules of 3PGA are phosphorylated by 6 molecules of ATP (produced in
the light reaction) to yield 6 molecules of 1-3 diphospho glyceric acid and 6 molecules of
ADP. This reaction is catalyzed by the enzyme, Kinase
3 Phospho + ATP
glyceric acid

Kinase

1,3 diphospho
glyceric acid

ADP

Six molecules of 1, 3 diphosphoglyceric acid are reduced with the use of 6 molecules of
NADPH2 (produced in light reaction) to form 6 molecules of 3 phospho glyceraldehyde.
This reaction is catalysed by the enzyme, triose phosphate dehydrogenase.
1,3 diphospho
glyceric acid

NADPH2

Triose phosphate
Dehydrogenase

3 phospho
glyceraldehyde

+ NADP +

H3PO4

3. Regenerative phase
In the regenerative phase, the ribose diphosphate is regenerated. The regenerative
phase is called as pentose phosphate pathway or hexose monophophate shunt. It involves
the following steps.
1. Some of the molecules of 3 phospho glyceraldehyde into dihydroxy acetone
phosphate. Both 3 phospho glyceraldehyde and dihydroxy acetone phosphate then
unite in the presence of the enzyme, aldolase to form fructose, 1-6 diphosphate.
3 phospho
glyceraldehyde

Triose phosphate isomerase

Dihydroxy acetone PO4

(DHAP)

3 phospho
Aldolase
Fructose 1,6 diphosphate
glyceraldehyde
+ DHAP
2. Fructose 6 phosphate is converted into fructose 6 phosphate in the presence of
phosphorylase
Fructose 1,6 diphosphate

Phosphorylase

Fructose 6 phosphate

3. Some of the molecules of 3 phospho glyceraldehyde instead of forming hexose

sugars are diverted to regenerate ribulose 1-5 diphosphate
3 phospho glyceraldehyde

Ribulose 1,5 diphosphate

4. 3 phospho glyceraldehyde reacts with fructose 6 phosphate in the presence of

enzyme transketolase to form erythrose 4 phosphate ( 4C sugar) and xylulose 5
phosphate(5C sugar)
3 phospho
glyceraldehyde

Fructose 6
+ phosphate

Transketolase
80

Erythrose 4 phosphate +
Xylulose 5 phosphate

5. Erythrose 4 phosphate combines with dihydroxy acetone phosphate in the

presence of the enzyme aldolase to form sedoheptulose 1,7 diphosphate(7C sugar)
Erythrose 4 phosphate
6.

+ DHAP

Aldolase

Sedoheptulose 1 ,7
diphosphate

Sed
oheptulose 1, 7 diphosphate loses one phosphate group in the presence of the
enzyme phosphatase to form sedoheptulose 7 phosphate.
Sedoheptulose 1 ,7 + ADP
diphosphate

Phosphatase

Sedoheptulose 7
phosphate

+ ATP

7. Sedoheptulose phosphate reacts with 3 phospho glyceraldehyde in the presence

of transketolase to form xylulose 5 phosphate and ribose 5 phosphate ( both % c
sugars)
Sedoheptulose
7 phosphate

3 phospho
glyceraldehyde

Transketolase

Xylulose
5 phosphate

Ribose 5
+ phosphate

8. Ribose 5 phosphate is converted into ribulose 1, 5 diphosphate in the presence of

enzyme, phosphopentose kinase and ATP. Two molecules of xylulose phosphate
are also converted into one molecule of ribulose monophosphate. The ribulose
monophosphate is phosphorylated by ATP to form ribulose diphosphate and ADP,
thus completing Calvin cycle.
Ribose
5 phosphate

ATP

Phophopentokinase

Ribulose 1,5 + ADP

diphosphate

2 mols of Xylulose Phophopentokinase

1 mol of Ribulose mono
Ribulose
+
ATP
Phophopentokinase
Ribulose
+ ADP
phosphate
phosphate
mono
diphosphate
phosphate
In the dark reaction, CO2 is fixed to carbohydrates and the CO2 acceptor ribulose
diphosphate is regenerated. In Calvin cycle, 12 NADPH 2 and 18 ATPs are required to fix
6 CO2 molecules into one hexose sugar molecule (fructose 6 phosphate).

81

6 CO2 +
12 NADPH2 +
18 ATP

Fructose 6 phosphate +

12 NADP+
18 ADP+
17 Pi

Schematic diagram of light reaction and Calvin cycle

C4 cycle or Hatch and Slack pathway

It is the alternate pathway of C3 cycle to fix CO 2. In this cycle, the first formed
stable compound is a 4 carbon compound viz., oxaloacetic acid. Hence it is called C4
cycle. The path way is also called as Hatch and Slack as they worked out the pathway in
1966 and it is also called as C4 dicarboxylic acid pathway. This pathway is commonly
seen in many grasses, sugar cane, maize, sorghum and amaranthus.
The C4 plants show a different type of leaf anatomy. The chloroplasts are
dimorphic in nature. In the leaves of these plants, the vascular bundles are surrounded by
bundle sheath of larger parenchymatous cells. These bundle sheath cells have
chloroplasts. These chloroplasts of bundle sheath are larger, lack grana and contain starch
grains. The chloroplasts in mesophyll cells are smaller and always contain grana. This
peculiar anatomy of leaves of C4 plants is called Kranz anatomy. The bundle sheath cells
are bigger and look like a ring or wreath. Kranz in German means wreath and hence it is
called Kranz anatomy. The C4 cycle involves two carboxylation reactions, one taking
place in chloroplasts of mesophyll cells and another in chloroplasts of bundle sheath
cells. There are four steps in Hatch and Slack cycle:
1. Carboxylation
2. Breakdown
3. Splitting
4. Phosphorylation
1. Carboxylation
82

It takes place in the chloroplasts of mesophyll cells. Phosphoenolpyruvate, a 3

carbon compound picks up CO2 and changes into 4 carbon oxaloacetate in the presence of
water. This reaction is catalysed by the enzyme, phosphoenol pyruvate carboxylase.
Phosphoenol
Pyruvate (3C)

CO2 +

H2O

PEP carboxylase

Oxaloacetate +
(4C)

H3PO4

2. Breakdown
Oxaloacetate breaks down readily into 4 carbon malate and aspartate in the
presence of the enzyme, transaminase and malate dehydrogenase.
Oxaloacetate (4C)

Transaminase

Aspartate (4C) +

Malate (4C)

Malate dehydrogenase

These compounds diffuse from the mesophyll cells into sheath cells.
3. Splitting
In the sheath cells, malate and aspartate split enzymatically to yield free CO 2 and
3 carbon pyruvate. The CO2 is used in Calvins cycle in the sheath cell.
Malate

Decarboxylation

CO2 + Pyruvate

The second Carboxylation occurs in the chloroplast of bundle sheath cells. The
CO2 is accepted by 5 carbon compound ribulose diphosphate in the presence of the
enzyme, carboxy dismutase and ultimately yields 3 phosphoglyceric acid. Some of the 3
phosphoglyceric acid is utilized in the formation of sugars and the rest regenerate
ribulose diphosphate.
4. Phosphorylation
The pyruvate molecule is transferred to chloroplasts of mesophyll cells where, it
is phosphorylated to regenerate phosphoenol pyruvate in the presence of ATP. This
reaction is catalysed by pyruvate phosphokinase and the phophoenol pyruvate is
regenerated.
Pyruvate

ATP +

Pi

Pyruvate
phosphokinase

Phosphoenol +
pyruvate

AMP +

Pyrophosphate

In Hatch and Slack pathway, the C3 and C4 cycles of carboxylation are linked and this is
due to the Kranz anatomy of the leaves. The C4 plants are more efficient in
photosynthesis than the C3 plants. The enzyme, phosphoenol pyruvate carboxylase of the
C4 cycle is found to have more affinity for CO2 than the ribulose diphosphate
carboxylase of the C3 cycle in fixing the molecular CO2 in organic compound during
Carboxylation.

83

Crassulacean Acid Metabolism (CAM) cycle or the dark fixation of CO2 in succulents
CAM is a cyclic reaction occurring in the dark phase of photosynthesis in the
plants of Crassulaceae. It is a CO2 fixation process wherein, the first product is malic acid.
It is the third alternate pathway of Calvin cycle, occurring in mesophyll cells. The plants
exhibiting CAM cycle are called CAM plants. Most of the CAM plants are succulents
e.g., Bryophyllum, Kalanchoe, Crassula, Sedium, Kleinia etc. It is also seen in certain
plants of Cactus e.g. Opuntia, Orchid and Pine apple families.
CAM plants are usually succulents and they grow under extremely xeric
conditions. In these plants, the leaves are succulent or fleshy. The mesophyll cells have
larger number of chloroplasts and the vascular bundles are not surrounded by well
defined bundle sheath cells.
In these plants, the stomata remain open during night and closed during day time. The
CAM plants are adapted to photosynthesis and survival under adverse xeric conditions.
CAM plants are not as efficient as C4 plants in photosynthesis. But they are better suited
to conditions of extreme desiccation.
CAM involves two steps:
1. Acidification
2. Deacidification
Acidification
In darkness, the stored carbohydrates are converted into phophoenol pyruvic acid
by the process of Glycolysis. The stomata in CAM plants are open in dark and they allow
free diffusion of CO2 from the atmosphere into the leaf. Now, the phosphoenolpyruvic
acid carboxylated by the enzyme phosphoenol pyruvic acid carboxylase and is converted
in to oxalaoacetic acid.
Phosphoenol Pyruvate

CO2 +

H2O

PEP carboxylase

84

Oxaloacetic acid +

H3PO4

The oxaloacetic acid is then reduced to malic acid in the presence of the enzyme
malic dehydrogenase. The reaction requires NADPH2 produced in Glycolysis.
Oxaloacetic acid

NADPH2 +

Malic dehydrogenase

Malic acid +

NADP+

The malic acid produced in dark is stored in the vacuole. The malic acid increases the
acidity of the tissues.
Deacidification
During day time, when the stomata are closed, the malic acid is decarboxylated to
produce pyruvic acid and evolve carbon dioxide in the presence of the malic enzyme.
When the malic acid is removed, the acidity decreases the cells. This is called
deacidification. One molecule of NADP+ is reduced in this reaction.
Malic acid

NADP+

Malic enzyme

Pyruvic acid +

NADPH2 +

CO2

The pyruvic acid may be oxidized to CO2 by the pathway of Krebs cycle or it
may be reconverted to phosphoenol pyruvic acid and synthesize sugar by C3 cycle. The
CO2 released by deacidification of malic acid is accepted by ribulose diphosphate and is
fixed to carbohydrate by C3 cycle.
CAM is a most significant pathway in succulent plants. The stomata are closed
during day time to avoid transpiration loss of water. As the stomata are closed, CO 2
cannot enter into the leaves from the atmosphere. However, they can carry out
photosynthesis during the day time with the help of CO 2 released from organic acids.
During night time, organic acids are synthesized in plenty with the help of CO 2 released
in respiration and the CO2 entering from the atmosphere through the open stomata. Thus,
the CO2 in dark acts as survival value to these plants.

Lecture No: 13

85

DIFFERENCES BETWEEN C3, C4 AND CAM PATHWAYS - FACTORS AFFECTING

PHOTOSYNTHESIS
Comparison of the plants of C3 and C4 cycle
C3 Plant

C4 Plant

1.

Only C3 cycle is found

Both C4 and C3 cycles are found.

2.

The efficiency of CO2 absorption at low

concentration is far less and hence, they
are less efficient.

The efficiency of CO2 absorption at

low concentration is quite high and
hence, they are more efficient plants.

3.

The CO2 acceptor is Ribulose-1, 5diphosphate.

The CO2 acceptor is phospho enol

pyruvate.

4.

The first stable product is phospho

glyceric acid (PGA).

Oxaloacetate (OAA) is the first stable

product.

5.

Plants show one type of chloroplast

(monomorphic type).

Plants show dimorphic type of

chloroplast. The chloroplast of
parenchymatous bundle sheath is
different from that of mesophyll cells
(dimorphic type). The chloroplasts in
bundle sheath cell are centripetally
arranged and lack grana. Leaves show
Kranz type of anatomy.

6.

In each chloroplast, two pigment

systems (Photosystem I and II) are
present.

In the chloroplasts of bundle sheath

cells, the photosystem II is absent.
Therefore, these are dependent on
mesophyll chloroplasts for the supply
of NADPH + H+.

7.

The Calvin cycle enzymes are present in

mesophyll chloroplast. Thus, the Calvin
cycle occurs.

Calvin cycle enzymes are absent in

mesophyll chloroplasts. The cycle
occurs only in the chloroplasts of
bundle sheath cells.

8.

The CO2 compensation point is 50-150

ppm CO2.

The CO2 compensation point is 0-10

ppm CO2.

9.

Photorespiration is present and easily

detectable.

Photorespiration is present only to a

slight degree or absent.

86

10. The CO2 concentration inside leaf

remains high (about 200 ppm).

The CO2 concentration inside the leaf

remains low (about 100 ppm).

11. The 13C/12C ratio in C-containing

compounds remains relatively low (both
13
CO2 and 12CO2 are present in air).

The ratio is relatively high, i.e. C4

plants are more enriched with 13C than
C3 plants.

12. Net rate of photosynthesis in full

It is 40-80 mg. of CO2 per dm2 of leaf
sunlight (10,000 12,000 ft. c.) is 15-25 area per hour. That is, photosynthetic
mg. of CO2 per dm2 of leaf area per hour. rate is quite high. The plants are
efficient.
13. The light saturation intensity reaches in
the range of 1000-4000 ft. c.

It is difficult to reach saturation even

in full sunlight.

14. Bundle sheath cells are unspecialized.

The bundle sheath cells are highly

developed with unusual construction
of organelles.

15. The optimum temperature for the

process is 10-25C.

In these plants, it is 30-45C and

hence, they are warm climate plants.
At this temperature, the rate of
photosynthesis is double than that is in
C3 plants.

16. 18 ATPs are required to synthesize one

glucose molecule.

30 ATPs are required to synthesize one

glucose molecule.

Factors affecting photosynthesis

I. External factors
1. Light
It is the most important factor of photosynthesis. Any kind of artificial light such
as electric light can induce photosynthesis. Out of the total solar energy, only 1-2 % is
used for photosynthesis and the rest is used for other metabolic activities. The effect of
light on photosynthesis can be studied under three categories.
a. Light intensity
Wolkoff (1966) found that the arte of photosynthesis is directly proportional to
light intensity. But the extremely high light intensities do not favor for higher
photosynthetic rates. The high light intensity which fails to accelerate photosynthesis is

87

called light saturation intensity. Of the light falling on a leaf, about 80 per cent is
absorbed, 10 per cent is reflected and 10 % is transmitted. The rate of photosynthesis is
greater in intense light than in diffused light. The plants are grouped into two types on the
basis of light requirement.
i. Heliophytes (Sun plants)
ii. Sciophytes (Shade plants)
At a specific light intensity, the amount of CO2 used in photosynthesis and the
amount of CO2 released in respiration are volumetrically equal. This specific light
intensity is known as light compensation point.
At very high light intensity, beyond a certain point, the photosynthetic cells
exhibit photo oxidation. This phenomenon is called solarisation and a result of this,
inactivation of chlorophyll molecules, bleaching of chlorophyll molecules and even
inactivation of some enzymes take place resulting in the destruction of whole
photosynthetic apparatus. In general, low light intensity favours stomatal closure and in
turn reduced rate of photosynthesis.
b. Light quality (wavelength)
Photosynthesis occurs only in the visible part of the light spectrum i.e., between
400 and 700 nm. The maximum rate of photosynthesis occurs at red light followed by
blue light. The green light has minimum effect and photosynthesis cannot take place
either in the infrared or in the ultraviolet light.
c. Light duration
In general tropical plants get 10-12 hours of light per day and this longer period of
light favours photosynthesis.
2. Carbon dioxide
CO2 is one of the raw materials required for photosynthesis. If the CO 2
concentration is increased at optimum temperature and light intensity, the rate of
photosynthesis increases. But, it is also reported that very high concentration of CO 2 is
toxic to plants inhibiting photosynthesis.

3. Temperature

88

The rate of photosynthesis increases by increase in temperature up to 40 C and

after this, there is reduction in photosynthesis. High temperature results in the
denaturation of enzymes and thus, the dark reaction is affected. The temperature
requirement for optimum photosynthesis varies with the plant species. For example,
photosynthesis stops in many plants at 0 C but in some conifers, it can occur even at -35
C. Similarly photosynthesis stops beyond 40-50 C in certain plants; but certain bacteria
and blue green algae can perform photosynthesis even at 70 C.
4. Water
Water has indirect effect on the rate of photosynthesis although it is one of the raw
materials for the process. The amount of water utilized in photosynthesis is quite small
and even less than 1 per cent of the water absorbed by a plant. Water rarely acts as a
limiting factor for photosynthesis. During water scarcity, the cells become flaccid and the
rate of photosynthesis might go down.
5. Oxygen
Oxygen is a byproduct of photosynthesis and an increase in the O 2 concentration
in many plants results in a decrease in the rate of photosynthesis. The phenomenon of
inhibition of photosynthesis by o2 was first discovered by Warburg (1920) in green alga
Chlorella and this effect is known as Warburgs effect. This is commonly observed in C3
plants.
In plants, there is a close relationship between Warburgs effect and
photorespiration. The substrate of photorespiration is glycolate and it is synthesized from
some intermediates of Calvins cycle. In plants that show Warburgs effect, increased O 2
concentration result in diversion of these intermediates of Calvin cycle into the synthesis
of glycolate, thereby showing higher rate of photorespiration and lower photosynthetic
productivity.
6. Mineral elements
The elements like Mg. Fe, Cu, Cl, Mn, P etc are involved in the key reactions of
photosynthesis and hence, the deficiency of any of these nutrients caused reduction in
photosynthesis.
7. Chlorophyll content

89

It is very much essential to tarp the light energy. In 1929, Emerson found direct
relationship between the chlorophyll content and rate of photosynthesis. In general, the
chlorophyll sufficient plants are green in colour showing efficient photosynthesis. The
chlorotic leaves due to irregular synthesis of chlorophyll or breakdown of chlorophyll
pigment exhibit inefficient photosynthesis.
8. Leaf
The leaf characters such as leaf size, chlorophyll content, number of stomata. Leaf
orientation and leaf age are some of the factors that are responsible for photosynthesis.
The maximum photosynthetic activity is usually seen in the physiologically functional
and full size leaves (usually third/fourth leaf from the tip of the shoot system).
9. Carbohydrates
If the accumulated carbohydrates are not translocated, the photosynthetic rate is
reduced and respiration is increased. Sugar is converted into starch and gets accumulated
in the chloroplasts. This reduces the effective surface in the chloroplast and the rate of
photosynthesis is decreased.
10. Phytohormones
Treharne (1970) reported first that photosynthesis may be regulated by plant
hormone system. He found that gibberellic acid and cytokinin increase the carboxylating
activity and photosynthetic rates. Meidner (1967) also reported that kinetin @ 3m
causes 12 per cent increase in photosynthesis within one hour of the treatment.

90

Lecture No: 14
PHOTORESPIRATION - PHOTORESPIRATION PROCESS AND SIGNIFICANCE OF
PHOTORESPIRATION
The excessive respiration that takes place in green cells in the presence of light is
called as photorespiration. Decker (1955) discovered the process and it is also called as
C2 cycle as the 2 carbon compound glycolic acid acts as the substrate in photorespiration.
In general, respiration takes place under both light and dark conditions. However in some
plants, the respiration is more in light than in dark. It is 3-5 times higher than the rate of
respiration in dark. Photorespiration is carried out only in the presence of light. But the
normal respiration is not light dependent and it is called dark respiration.
In photorespiration, temperature and oxygen concentration play an important role.
Photorespiration is very high when the temperature is between 25 and 30 C. The rate of
photorespiration increases with the increase in the concentration of oxygen. Three cell
organelles namely chloroplast, peroxisome and mitochondria are involved in the
photorespiration. This kind of respiration is seen in plants like cotton, pulses, capsicum,
peas, tomato, petunia soybean, wheat, oats, paddy, chlorella etc and it is absent in
grasses.
Mechanism
1. In the presence of excess oxygen and low CO 2 , ribulose 1,5 diphosphate produced in
the chloroplast during photosynthesis is split into 2 phospho glycolic acid and 3 phospho
glyceric acid by the enzyme, ribulose 1,5 diphosphate oxygenase
2. The 3 phospho glyceric acid enters the Calvin cycle.
3. In the next step, phosphate group is removed from 2 phosphoglycolic acid to produce
glycolic acid by the enzyme, phosphatase.
4. Glycolic acid then it come out of chloroplast and enter the peroxisome. Here, it
combines with oxygen to form glyoxylic acid and hydrogen peroxide. This reaction is
catalyzed by the enzyme, glycolic acid oxidase. Hydrogen peroxide is toxic and it is
broken down into water and oxygen by the enzyme, Catalase. Photorespiration is an
oxidation process. In this process, glycolic acid is converted into carbohydrate and CO 2 is
released as the by product. As glycolic acid is oxidized in photorespiration, it is also
called as glycolate metabolism.

91

5. The glyoxylic acid converted into glycine by the addition of one amino group with the
help of the enzyme, amino transferase.
6. Now, the glycine is transported from the peroxisome into the mitochondria. In the
mitochondria, two molecules of glycine condense to form serine and liberate carbon
dioxide and ammonia.
7. Amino group is removed from serine to form hydroxyl pyruvic acid in the presence of
the enzyme, transaminase.
8. Hydroxy pyruvic acid undergoes reduction with the help of NADH to form glyceric
acid in the presence of enzyme alpha hydroxyl acid reductase.
9. Finally, regeneration of 3 phosphoglyceric acid occurs by the phosphorylation of
glyceric acid with ATP. This reaction is catalyzed by the enzyme, Kinase.
10. The 3 phosphoglyceric acid is an intermediate product of Calvin cycle. If it enters the
chloroplast, it is converted into carbohydrate by photosynthesis and it is suppressed
nowadays with the increased CO2 content in the atmosphere.

Significance of photorespiration
1.Photorespiration helps in classifying the plants
Generally, photorespiration is found in C3 plants and absent in C4 plants.
2. Carbon dioxide is evolved during the process and it prevents the total depletion of CO 2
in the vicinity of chloroplasts.
92

2. The process causes oxidation of glycolic acid which arises as an unwanted byproduct
of photosynthesis. The glycolic acid after oxidation is converted into carbohydrate but the
remainder is converted into CO2.
3. Photorespiration uses energy in the form of ATP and reduced nucleotides, but normal
respiration yields ATP and reduced nucleotides.
4. It is believed that photorespiration was common in earlier days when CO 2 content was
too low to allow higher rates.

93

Lecture No: 15
RESPIRATION - GLYCOLYSIS, TCA AND PENTOSE PHOSPHATE
PATHWAY
The cellular oxidation or break down of carbohydrates into CO 2 and H2O, and
release of energy is called as respiration. It is a reverse process of photosynthesis. In
respiration, the oxidation of various organic food substances like carbohydrates, fats,
proteins etc, may take place. Among these, glucose is the commonest.
C6H12O6 + 6O2 6CO2 + 6H2O + Energy (686 kcal)
This oxidation process in not so simple and does not take place in one step.
Breakdown of glucose involves many steps releasing energy in the form of ATP
molecules and also forming a number of carbon compounds (intermediates). Respiration
is a vital process that occurs in all living cells of the plant and the most actively respiring
regions are floral buds, vegetative buds, germinating seedlings, stem and root apices.
Types of respiration
Degradation of organic food for the purpose of releasing energy can occur with or
without the participation of oxygen. Hence, respiration can be classified into two types;
aerobic and anaerobic respiration.
Aerobic respiration
Aerobic respiration takes place in the presence of oxygen and the respiratory
substrate gets completely oxidized to carbon dioxide and water as end products.
C6H12O6 + 6O2 6CO2 + 6H2O + Energy (686 kcal)
(Glucose)
This type of respiration is of common occurrence and it is often used as a synonym of
respiration.
Anaerobic respiration
It takes place in the absence of oxygen and the respiratory substrate is
incompletely oxidized. Some other compounds are also formed in addition to carbon

94

dioxide. This type of respiration is of rare occurrence but, common among

microorganisms like yeasts.
C6H12O6 2C2 H5OH + 2CO2 + 56 kcal
Glucose

Ethanol

Respiratory substrate
A respiratory substrate is an organic substance which can be degraded to produce
energy which is required for various activities of the cell. The respiratory substrates
include carbohydrates, fats, organic acids, protein etc.
Carbohydrates
The carbohydrates constitute the most important respiratory substrate and the
common amongst them are starch, sucrose, glucose and fructose. The complex
carbohydrates are first hydrolyzed to simple sugars and then they are utilized.
Starch Disaccharides Hexoses
Fats
The fats are important storage food in seeds. Nearly 80 per cent of the
angiosperms have fats as the main storage food in their seeds. At the time of seed
germination, large amount of fats are converted into carbohydrates while the remaining
fats are utilized in respiration. Fats are first broken down to glycerol and fatty acids. The
fatty acids are broken down to acetyl coenzyme by -oxidation. The acetyl coenzyme
enters Krebs cycle for further degradation and releases energy. Glycerol can directly
enter the respiratory channel via glyceraldehyde.
Organic acids
Organic acids normally do not accumulate in plants to any appreciable extent
except in the members of the family, Crassulaceae. Organic acids are oxidized under
aerobic conditions to carbon dioxide and water.
Proteins
Under normal conditions, proteins are used up as respiratory substrate only in
seeds rich in storage proteins. In vegetative tissues, proteins are consumed only under
starvation. The proteins are hydrolyzed to form amino acids. Later, the amino acids

95

undergo deamination forming organic acids and the organic acids can enter Krebs cycle
directly.
Mechanism of Respiration
1. Glycolysis
2. Aerobic breakdown of pyruvic acid (Krebs cycle)
3. Electron Transport System/ Terminal oxidation / oxidative phosphorylation
5. Pentose phosphate pathway
A. GLYCOLYSIS / EMBDEN MEYER HOF PARANAS (EMP) PATHWAY
Glycolysis can take place even in the absence of O2. One molecule of the 6 carbon
compound, glucose is broken down through a series of enzyme reactions into two 3carbon compounds, the pyruvic acid. Glycolysis takes place in the cytoplasm and it does
not require oxygen. Hence it is an anaerobic process.
1. Glucose molecules react with ATP molecules in the presence of the enzyme
hexokinase to form glucose -6- phosphate.
Glucose + ATP Glucose -6- phosphate + ADP
2. Glucose-6-phosphate is isomerised into fructose-6-phosphate in the presence of
phospho hexose isomerase.
Fructose + ATP Fructose -6- phosphate + ADP
3. Fructose-6-phosphate reacts with one molecule of ATP in the presence of phospho
hexo kinase forming fructose 1, 6-disphosphate.
Fructose 6- phosphate + ATP Fructose -1,6- biphosphate + ADP
4. Fructose 1, 6 diphosphate is converted into two trioses, 3-phospho glyceraldehyde
and dihydroxy acetone phosphate in the presence of aldolase.
Fructose -1,6- biphosphate 3-phospho glyceraldehyde+ DHAP
5. 3-phosphoglyceraldehyde

reacts

with

H3PO4

and

diphosphoglyceraldehyde where, the reaction is non enzymatic.

96

forms

1,3-

6. 1, 3-Diphosphoglyceraldehyde is oxidized to form 1,3- diphosphoglycerate in the

presence of triose-phosphate dehydrogenase and coenzyme NAD+. The NAD+ acts as
hydrogen acceptor and reduced to NADH+ + H+ in the reaction.
Glyceraldehde -3- phosphate + NAD + Pi 1,3- diphosphoglycerate + NADH
6. 1, 3-Diphosphoglycerate reacts with ADP in the presence of phosphoglyceric
transphorylase (kinase) to form 3 phosphoglyceric acid and ATP.
1,3- diphosphoglycerate + ADP 3, Phosphoglycerate + ATP
7. 3, Phosphoglycerate

2, Phosphoglycerate acid is isomerized into 2

phosphoglyceric acid in the presence of the enzyme, phospho glycero mutase

3, Phosphoglycerate 2, Phosphoglycerate
8. 2 phosphoglyceric acid is converted into 2-phosphoenolpyruvic acid in the
presence of enolase.
2, Phosphoglycerate Phosphoenol pyruvate + H2O
9. 2 phospho enol pyruvic acid reacts with ADP to form one molecule each of
pyruvic acid and ATP in the presence of pyruvate kinase.
Phosphoenol pyruvate + ADP Pyruvate + ATP
Glycolysis or EMP pathway is common in both aerobic and anaerobic respiration.
The overall glycolytic process can be summarized as follows.
C6H12O6 + 2ATP + 2NAD + 4ADP+2H3PO4

2 CH3COCOOH + 2ADP + 2NADH2 + 4 ATP

Pyruvic acid

Thus there is a gain of 4-2 = 2 ATP molecules per hexose sugar molecule oxidized
during this process.

97

Besides this, 2 molecules of reduced coenzyme NADH 2 are also produced per
molecule of hexose sugar in glycolysis.

During aerobic respiration, these two NADH2 are oxidized via the electron
transport chain to yield 3 ATP molecules each. Thus 6 ATP molecules are formed.

98

99

B. KREBS CYCLE / CITRIC ACID CYCLE /TCA CYCLE

The pyruvic acid produced in glycolysis enters into Krebs cycle for further
oxidation. Krebs cycle is also known as citric acid cycle or Tri carboxylic acid (TCA)
cycle. This aerobic process takes place in mitochondria where necessary enzymes are
present in matrix.
11. Pyruvic acid reacts with CoA and NAD and is oxidatively decarboxylated. One
molecule of CO2 is released and NAD is reduced. Pyruvic acid is converted into acetyl
CoA.
Pyruvate dehydrogenase
Acetyl COA + CO2 + NADH2
Pyruvic acid + CoA + NAD
12. Acetyl-CoA condenses with oxaloacetic acid in the presence of condensing enzyme
and water molecule to form citric acid. CoA becomes free.
Condensing enzyme
Citric acid + CoA
Acetyl CoA + Oxaloacetic acid
+ H2O
13. Citric acid is dehydrated in the presence of aconitase to form cis aconitic acid
Aconitase
Cis Aconitic acid

Citricacid
- H2O

14. Cis-aconitic acid reacts with one molecule of water to form Isocitric acid
Isocitric acid

Cis-aconitic acid + H2O

15. Iso-citric acid is oxidized to oxalo succinic acid in the presence of Isocitric
dehydrogenase. NADP is reduced to NADPH2 in the reaction.
IC dehydrogenase
Oxalo succinic acid + NADPH2

Isocitric acid + NADP

100

16. Oxalo succinic acid is decarboxylated in the presence of oxalo succinic decarboxylase
to form - ketoglutaric acid and a second molecule of CO2 is released.
Oxalosuccinic
Oxalosuccinic acid

-ketoglutaric acid + CO2

Decarboxylase
17. - ketoglutaric acid reacts with CoA and NAD in the presence of - ketoglutaric
acid dehydrogenase complex and is oxidatively decarboxylated to form succinyl CoA and
a third mole of CO2 is released. NAD is reduced in the reaction.
Succinyl-CoA + CO2 + NADH2

-keto glutaric acid + CoA

NAD NADH2
18. Succinyl CoA reacts with water molecule to form succinic acid. CoA becomes free
and one molecule of GDP (Guanosine diphosphate) is phosphorylated in presence of
inorganic phosphate to form one molecule of GTP.
H2O
Succinic acid + GTP
Succinyl-CoA + GDP + ip
GTP may react with ADP to form one molecule of ATP
GTP + ADP ATP + GDP
19. Succinic acid is oxidized to fumaric acid in the presence of succinic dehydrogenase
and co enzyme FAD is reduced in this reaction.
Succinic acid dehydrogenase
Fumaric acid + FADH2
Succinic acid + FAD
20. One mole of H2O is added to Fumaric acid in the presence of fumarase to form malic
acid.
Fumarase
Malic acid

Fumaric acid + H2O

21. In the last step, malic acid is oxidized to oxaloacetic acid in the presence of malic
dehydrogenase and one molecule of coenzyme i.e. NAD is reduced.

101

Malic dehydrogenase
Oxaloacetic acid + NADH2

Malic acid + NAD

KREBS CYCLE or TCA CYCLE

102

103

Pentose phosphate pathway (ppp) / hexose mono phosphate (hmp) shunt/

phosphogluconate pathway / warburg and dickens pathway
The pentose phosphate pathway occurs in the cytoplasm outside the mitochondria
and it is an alternative pathway to glycolysis and Krebs cycle. The presence of some
compounds like iodoacetate, fluorides, arsenates etc. inhibit some steps in glycolysis and
that leads to the alternate pathway. This pathway was discovered by Warburg and Dicken
(1938). This pathway does not produce ATP but it produces another form of energy called
reducing power in the form of NADPH. It is not oxidized in the electron transport system
but, it serves as hydrogen and electron donor in the biosynthesis of fatty acids and
steroids. The pentose phosphate pathway consists of two distinct phases. In the first
phase, hexose is converted into pentose and in the second phase, pentose is reconverted in
to hexose.
In the process, oxidation of glucose 6 phosphate leads to the formation of 6
phosphogluconic acid (pentose phosphate). Since glucose is directly oxidized without
entering glycolysis, it is called as direct oxidation.
6 Glucose 6 phosphate +12 NADP

5 Glucose 6 Phosphate + 12 NADPH2+ 6 CO2

It provides ribose sugars for the synthesis of nucleic acids and is also required for
shikimic acid pathway. Although ATP is not produced, NADPH is produced and serves as
hydrogen and electron donor in the biosynthesis of fatty acids and steroids. The pathway
is also called as phosphogluconate pathway as the first product in this pathway is
phosphogluconate.

104

Lecture No: 16
OXIDATIVE PHOSPHORYLATION
DIFFERENCES BETWEEN OXIDATIVE PHOSPHORYLATION AND
PHOTOPHOSPHORYLATION. RESPIRATORY QUOTIENT AND ENERGY
BUDGETING IN RESPIRATION.
C. TERMINAL OXIDATION OF THE REDUCED COENZYMES / ELECTRON
TRANSPORT SYSTEM AND OXIDATIVE PHOSPHORYLATION
The last step in aerobic respiration is the oxidation of reduced coenzymes
produced in glycolysis and Krebs cycle by molecular oxygen through FAD, UQ
(ubiquinone), cytochrome b, cytochrome c, cytochrome a and cytochrome a 3 (cytochrome
oxidase).
Two hydrogen atoms or electrons from the reduced coenzyme (NADH2 or
NADPH2) travel through FAD and the cytochromes and ultimately combines with 1/2O 2
molecule to produce one molecule of H2O. This is called as terminal oxidation.
The terminal oxidation of each reduced coenzyme requires 1/2O2 molecule and
2H atoms (i.e. 2 e- + 2H+) to produce one H2O molecule. Except for flavoproteins (like
FAD) and ubiquinone (UQ) which are hydrogen carriers, the other components of
electron transport chain (cytochromes) are only electron carriers i.e. they cannot give or
take protons (H+)
During the electron transport, FAD and the iron atom of different cytochromes get
successively reduced (Fe++) and oxidized (Fe+++) and enough energy is released in some
places which is utilized in the photophosphorylation of ADP molecules in the presence of
inorganic phosphate to generate energy rich ATP molecules. Since, this oxidation
accompanies phosphorylation; it is called as oxidative phosphorylation.
One molecule of ATP with 7.6 Kcal.energy is synthesized at each place when
electrons are transferred from
Reduced NADH2 or NADPH2 to FAD
1.
Reduced cytochrome b to cytochrome c
2.
Reduced cytochrome a to cytochrome a3
3.
Thus, oxidation of one molecule of reduced NADH 2 or NADPH2 will result in the
formation of 3 ATP molecules while the oxidation of FADH 2 lead to the synthesis of 2
ATP molecules.
According to the most recent findings, although in eukaryotes terminal oxidation
of mitochondrial NADH / NADPH results in the production of 3 ATP molecules but that
of extra mitochondrial NADH / NADPH yields only 2 ATP molecules. Therefore, the
two reduced coenzyme molecules (NADH) produced per hexose sugar molecule during
Glycolysis will yield only 2x2:4 ATP molecules instead of 6 ATP molecules. Complete
oxidation of a glucose molecule (hexose sugar) in aerobic respiration results in the net
gain of 36 ATP molecules in most eukaryotes.
One glucose molecule contains about 686 Kcal. Energy and 36 ATP molecules
will have 273.6 Kcal energy. Therefore about 40% (273.6/686) energy of the glucose
molecule is utilized during aerobic breakdown and the rest is lost as heat. Since huge
amount of energy is generated in mitochondria in the form of ATP molecules, they are
called as Power Houses of the cell.

105

ATP molecules contain energy in terminal pyrophosphate bonds. When these

energy rich bonds break, energy is released and utilized in driving various other
metabolic processes of the cell.
Differences between oxidative phosphorylation and Photophosphorylation
Oxidative phosphorylation
Photophosphorylation
1
It occurs during respiration
Occurs during photosynthesis
2
Occurs inside the mitochondria (inner
Occurs inside the chloroplast (in the
membrane of cristae)
thylakoid membrane)
3
Molecular O2 is required for terminal
Molecular O2 is not required
oxidation
4
Pigment systems are not involved
Pigment systems, PSI and PSII are
involved
5
It occurs in electron transport system
Occurs during cyclic and non cyclic
electron transport
6
ATP molecules are released to
ATP molecules produced are utilized
cytoplasm and used in various metabolic for CO2 assimilation in the dark
reactions of the cell
reaction of photosynthesis
Efficiency of respiration
The total energy content of one molecule of glucose is 686 Kcal. Out of this
energy, available free energy is 673.6 Kcal and the energy content of ATP molecule is
calculated as 7.3 Kcal. The efficiency of respiration may be expressed as follows.
Kcal of energy conserved in ATP
Efficiency of respiration: ---------------------------------------- x 100
Total free energy available
38 x 7.3
: ------------x 100: 41 %
673.6

Efficiency of aerobic respiration

2 x 7.3
Efficiency of anaerobic respiration: ------------x 100: 31 %
47
2 x 7.3
: ------------x 100: 36.5 %
40

Efficiency of fermentation

Respiratory quotient
The ratio of the volume of CO 2 released to the volume of O 2 taken during
respiration is called as respiratory quotient and is denoted as RQ
RQ

Volume of CO2
Volume of O2

106

Value of RQ
The value of RQ depends upon the nature of the respiratory substrate and the
amount of O2 present in respiratory substrate.
1. When carbohydrates such as hexose sugars are oxidized in respiration, the value of RQ
is 1 or unity because volume of CO2 evolved equals to the volume of O2 absorbed.
C6H12O6 + 6O2
6CO2 + 6H2O
Glucose
volume of CO2
6
RQ =
=
= 1 or unity
volume of O2
6
2. When fats are the respiratory substrate, the value of RQ becomes less than one because
fats are poorer in O2 in comparison to carbon and they require more O2 for their
oxidation,
2C51H98O6 +145O2
102CO2 + 98H2O
Tripalmitin
volume of CO2
102
RQ =
=
= 0.7
volume of O2
145
(Fats are oxidized in respiration usually during the germination of fatty seeds).
3. When organic acids are oxidized in respiration, the value of RQ becomes more than
one. It is because organic acids are rich in O2 and require less O2 for their oxidation.
C4H6O5 + 3O2
4CO2 + 3H2O
Malic acid
volume of CO2
4
RQ =
=
= 1.3
volume of O2
3
Energy budgeting
Stages

Gain of
ATP

Glycolysis
1) Glucose
Glucose 6 PO4
2) Fructose 6 PO4
Fructose 1,6 di PO4
3) 1,3 diphosphoglyceraldehyde
1,3 diphospho glyceric acid
4) 1,3 diphospho glyceric acid
3 phosphoglyceric acid
5) 2 phosphoenol pyruvic acid
Pyruvic acid
Total
Krebs cycle
6) Pyruvic acid
Acetyl CoA
7) Isocitric acid
Oxalosuccinic acid
8) ketoglutaric acid
Succinyl CoA
9) Succinyl Co A
Succinic Acid
10) Succinic acid
Fumaric Acid
11) Malic acid
Oxaloacetic acid
Total ATP mol. produced per Pyruvic acid
Total ATP mol. produced for 2 Pyruvic acids
Grand Total

107

Consumption
of ATP

Net gain of
ATP

1
1
6
2
2
10
3
3
3
1
2
3
15
15 x 2:30
40

-2

-2

15
30
8+ 30 = 38

108

LECTURE NO: 17
MID SEMESTER EXAMINATION

109

Lecture No: 18
FACTORS AFFECTING RESPIRATION - DIFFERENCE BETWEEN
PHOTORESPIRATION AND DARK RESPIRATION - ROLE OF
RESPIRATION

Factors affecting respiration

A. External factors
1. Temperature
Temperature has profound influence on the rate of respiration.

Optimum

temperature for respiration is about 30C, minimum 0C and maximum about 45C. At
low temperature, the respiratory enzymes becomes inactive, consequently the rate of
respiration falls. It is due to this fact that the quality of fruits and vegetables stored at low
temperature does not deteriorate. At very high temperature, respiration slows down and
may even be stopped due to denaturation of the respiratory enzymes.
2. Oxygen
In complete absence of O2, anaerobic respiration takes place while aerobic
respiration stops. In higher plants, the anaerobiosis produces large amount of alcohol
which is toxic to plants. If some amount of O2 is available, anaerobic respiration slows
down and aerobic respiration starts. The concentration of O2 at which aerobic respiration
is optimum and anaerobic respiration is stopped, is called as extinction point.
It is observed that under anaerobic conditions, much more sugar is taken up per
quantity of yeast present than it is consumed in the presence of oxygen. The inhibition on
the rate of carbohydrate breakdown by oxygen is called as Pasteurs effect.
4.

Carbon dioxide

5.

Higher concentration of CO2 in the atmosphere especially in the poorly aerated

soil has retarding effect on the rate of respiration.
4. Inorganic salts
If a plant or tissue is transferred from water to salt solution, the rate of respiration
increases (called as salt respiration).

110

5. Water
Proper hydration of cells is essential for respiration. Rate of respiration decreases
with decreased amount of water, so much so, that in dry seeds, the respiration is at its
minimum. It is because in the absence of a medium, the respiratory enzymes become
inactive.
6. Light
The effect of light is indirect on the rate of respiration through the synthesis of
organic food matter in photosynthesis.
7. Wound or injury
Injury or wounds result in increased respiration as the plants in such a state
require more energy which comes from respiration. The wounded cells become more
meristematic to form new cells for healing the wound.
Internal factors
1. Protoplasmic factors
The amount of protoplasm in the cell and its state of activity influence the rate of
respiration.

The rate of respiration is higher in young meristematic cells which divide actively
and requires more energy. Such cells have greater amount of protoplasm and no
vacuoles.

In old mature tissues, the rate of respiration is lower because of lesser amount of
active protoplasm

2. Concentration of respiratory substrate

Increased concentration of respirable food material brings about an increase in the
rate of respiration.
Under starvation conditions, such as in etiolated leaves, the rate of respiration
slows down considerably. If such etiolated leaves are supplied with sucrose solution for
few days even in dark conditions, the rate of respiration increases.

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Differences between Photorespiration and Dark respiration

1

Photorespiration
It occurs in the presence of light

Dark /Mitochondrial respiration

It occurs in the presence of both light and

The substrate is glycolate

dark.
The respiratory substrate may be

It occurs in chloroplast, peroxisome and

carbohydrate, fat or protein.

The process occurs in the cytoplasm and

mitochondria
It occurs in temperate plants like, wheat

mitochondria
It occurs in C4 plants ( maize and sugar

and cotton ( mainly in C3 plants)

It occurs in the green tissues of plants

cane)
It occurs in all the living plants( both

6
7

The optimum temperature is 25- 35C

This process increases with increased

green and non green )

It is not temperature sensitive
This process saturate at 2-3 percent O2 in

CO2 concentration.

the atmosphere and beyond this

Hydrogen peroxide is formed during the

concentration there is no increase.

Hydrogen peroxide is not formed.

reaction
9 ATP molecules are not produced,
10 Reduced coenzymes such as NADPH2,

Several ATP molecules are produced.

Reduced coenzymes such as NADPH2,

NADH2 and FADH2 are not produced.

11 One molecule of ammonia is released

NADH2 and FADH2 are produced.

No ammonia is produced

per molecule of CO2 released.

12 Phosphorylation does not occur

Oxidative phosphorylation occurs.

Differences between respiration and photosynthesis

1

Respiration
It is catabolic process resulting in the

Photosynthesis
It is an anabolic process resulting in the

2
3
4

destruction of stored food

Light is not essential for the process
Oxygen is absorbed in the process
Carbon dioxide and water are

manufacture of food.
Light is very much essential
Oxygen is liberated
Carbon dioxide is fixed to form carbon

produced
Potential energy is converted into

containing compound
Light energy is converted into chemical

Kinetic energy
Glucose and oxygen are the raw

energy (potential energy)

Carbon dioxide and water are the raw

112

materials
Energy is released during respiration

materials
Energy is stored during photosynthesis

8
9

and hence it is an exothermic process.

Reduction in the dry weight
Chlorophyllous tissues are not

and hence it is an endothermic process

Gain in the dry weight
Chlorophyllous tissues are essential for

necessary

the process

Differences between aerobic respiration and fermentation

Aerobic respiration
1 It occurs in all living cells of the plants

Fermentation
Occurs outside the plant cells and in

throughout the day and night

2 It takes place in the presence of oxygen
3 The end products are CO2 and H2O

certain microorganisms
Absence of oxygen
End products are CO2 and alcohol or other

4 It is not toxic to plants

5 Complete oxidation is food material is

organic acids
It is toxic to plants
Incomplete oxidation is observed

observed
6 Large amount of energy (673 kCal) is

Very small amount of energy (21 kCal) is

released per glucose molecule

7 The complete oxidation yields 38 ATP

released per glucose molecule

The incomplete oxidation in fermentation

molecules
yields only two ATP molecules
8 The enzyme, zymase is not required but Zymase is required in the case of
many other enzymes and coenzymes

carbohydrates

are required

113

Lecture No: 19
PROTEIN AND FAT SYNTHESIS.
Biosynthesis of protein
Protein is a complex organic nitrogenous substance found in all living tissues of
plants and animals. They are polymer of amino acids in linear order. Synthesis of
protein may take place from amino acids produced by direct amination of organic acids
or by degradation of protein. Former is known as primary protein synthesis while the
latter is called secondary protein synthesis. Protein synthesis occurs in pre DNA
synthesis phase (G1 phase) of cell cycle.
Biosynthesis of protein takes place in prokaryotes as well as in eukaryotes. Kinds
of protein to be synthesized depend upon the gene (DNA segment). Gene is continuous
uninterrupted sequence of nucleotides which codes for a single polypeptide chain. Now
it is believed that the sequence of some eukaryotic genes is found to be interrupted by
nucleotides that are not represented with the amino acid sequence of protein. They are
non-coding (silent).

Genes control all metabolic processes by synthesizing proteins

(enzymes).
Structure of an eukaryotic gene showing Exon (coding part) and Intron (non-coding part).
Mechanism of protein synthesis: Protein synthesis takes place in following two stages:
I. Transcription
II. Translation
Transcription: Transcription occurs throughout inter phase and continues up to early
prophase of cell division. It is primary stage of protein synthesis. When DNA produces
DNA the process is called replication but when DNA produces RNA the process is called
transcription. In the former case DNA is duplicated while in the latter case protein is
synthesized.

During transcription RNA is synthesized on DNA template.

114

Here,

information or order contained in DNA is passed on the mRNA for synthesis of particular
protein. The information is in coded form and consists of three nitrogenous bases (triplet
codons). The part of DNA responsible for synthesis of mRNA is which leads to one
polypeptide chain is called cistron (functional gene).

New strands of mRNA are

synthesized on DNA template making use of RNA nucleotides present in surroundings in

5 3 director just like DNA chains.
Single DNA strand serves as template for RNA polymerase and synthesizes RNA.
DNA strand which serves as template for transcription is called the sense strand. The
complementary strand is antisense strand. In SV 40 viruses both strands of DNA are
transcribed and it is called symmetrical transcription, but when only one strand is
transcribed it is called asymmetrical transcription. Former type of transcription also
occurs in Polymer virus DNA and in the mitochondria genome.
Mechanism of transcription
Transcription involves following events:
I. Uncoiling of DNA molecule
II. Synthesis and action of enzyme RNA polymerase
III. Synthesis of hn RNA / mRAN.
Uncoiling of DNA molecule:

As per nucleosome model of chromosome.

Chromosomes are matarmala like beaded structure. Beads are separated by string and
are making up of DNA and histone protein. Histone proteins are of 5 kinds: H2A, H2B,
H3, H4 and H5 or H1. First four constitute the bead core and H1 links two beads. The
DNA molecules are wrapped on histone protein cores and linker protein core in beaded
and linker regions respectively.
Structure and function of enzyme RNA polymerase:

RNA polymerase is a

holoenzyme. Core particle consists of sub units , and . Cofactor consists of sigma
factor. For functional RNA polymerase formation the two (core enzyme and sigma
factor) gene united. Sigma factor recognized correct start signal at DNA template and

115

core enzyme continues transcription.

Sigma factor dissociates after initiation of

transcription to adjure with other core enzyme of RNA polymerase. In prokaryotes RNA
polymerase is only one type while in eukaryotes.
Production of mRNA / hnRNA: In prokaryotes where nucleus is not well
organized mRNA is the direct product of transcription, while in eukaryotes the direct
product of transcription is hnRNA (heteronucleic RNA) and mRNA is derived from
hnRNA by cutting and splicing. HnRNA has coding and non-coding sequences. Coding
sequences are interrupted by non-coding sequences. Non coding sequences are removed
by splicing (cutting) by endonuclease enzyme and coding sequences are ligased together
to from mRNA. The spliced non-coding sequences are degraded within nucleus. It never
goes out of nucleus. Thus, only fraction of hnRNA is translocated to cytoplasm from
nucleus via nuclear pore.
In eukaryotes migration of mRNA from nucleus to cytoplasm via nuclear pore
occurs through poly a tail. According to another view, ribosomes pull the mRNA from
nucleus to cytoplasm. Now mRNA gets established in cytoplasm.
Translation: Translation is a process in which order (message) given by DNA to mRNA
for synthesis of particular protein is implemented (conveyed).

Genetic information

concealed in mRNA directs the synthesis of particular protein. These orders are in coded
form. This coded information (expressed through codons) is recognized by tRNA having
anticodons.

Anticodons are opposite to codons (codons and anticodons are

complementary to each other).

Mechanism of translation: It involves following events:
I. Activation and selection of amino acids
II. Transfer of amino acids to tRNA molecules
III. Formation of protein synthesizing apparatus and chain initiation
IV. Chain elongation
V. Chain termination

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FAT SYNTHESIS
Fat synthesis can be studied under the following heads: 1. Fat synthesis of Glycerol
There may be different methods of the formation of glycerol in plants, but one of
the very common methods is from dihydroxy acetone phosphate which is an intermediate
of glycolysis. Dihydroxyacetone phosphate is first reduced to glycerophosphate by the
enzyme glycerol 3 phosphate dehydrogenase. Co-enzyme NADH 2 is oxidized in this
reaction. glycerophosphate is then hydrolysed by glycerol phosphatase to liberate
phosphoric acid and forming glycerol.
2. Synthesis of Fatty acids
Long chain saturated fatty acids* are synthesized in plants from active two carbon
units, the acetyl CoA (CH3CO.CoA). Although the reactions of oxidation of fatty
acids are reversible, the fatty acids are not formed simply by the reverse reactions of
oxidation. Synthesis of fatty acids from CH3CO.CoA takes place step by step. In each
step the fatty acid chain is increased by two carbon atoms. Each step involves two
reactions
(i) In the first reaction which takes place in the presence of acetyl CoA carboxylase,
acetyl CoA combines with CO2 to form malonyl CoA (malonic acid is 3 C
compound). ATP provides energy while Mn++ and biotin are required as co-factors.
(ii) Malonyl CoA reacts with another molecule of CH 3CO.CoA in the presence of fatty
acid synthetase and Coenzyme NADPH2 to form Coenzyme A derivative butric acid
(butyric acid contains 4 atoms). One mol. Of CO 2, H2O and CoA are released while
NADPH2 oxidised in the reaction.
Butyryl CoA, in the next step will combine with malonyl CoA to form CoA derivative of
fatty acid containing 6-C atoms. This process is repeated till Coenzymes-A derivative of
long chain fatty acid (which may contain up to 16-18C atoms) is produced.

117

(As a matter of fact the enzyme fatty acid synthetase is not simple but a complex of many
enzymes (multienzyme complex) and an acyl carrier protein called as ACP**. And
actually the reaction (ii) described above only summarises a number of reactions involved
in the synthesis of fatty acid from acetyl CoA which can be grouped under 3
categories :
(a) Initiation reaction: In this reaction acetyl CoA transfers its acetyl group to one of
the SH groups of multienzyme complex i.e. fatty acid synthatase.
Unsaturated fatty acids are synthesized by denaturation of saturated fatty acid.
ACP is similar to CoA in having phosphopantetheine as the functional unit in their
structures. In CoA, it is esterified to Adenosine 3, 5 bisphosphate but in ACP, it is
esterified to serine of a protein chain consisting of 81 amino acids.
Chain elongation reactions: Six different reactions involved
(b) Chain elongation reactions
Six different reactions involved here are (i) malonyl transfer, (ii) condensation,
(iii) reduction, (iv) dehydration, (v) reduction and (vi) acyl transfer. Chain elongation
starts with the transfer of malonyl group from malonyl CoA to second SH group of
the multienzyme complex. Then, there is a condensation of the latter so that a 4 C unit
is produced. This unit by next three reactions i.e. reduction, dehydration and reduction is
converted into saturated 4 C unit (i.e. butyryl CoA). In acyl transfer reaction the fatty
acid residue is transferred back to the SH group to which the acetyl group was
transferred in initiation reaction. The cycle is repeated again and again with malonyl
transfer, condensation etc. till the fatty acid residue consists of up to 16 - 18 C atoms.
Each such turn elongates fatty acid chain by 2 C atoms. Details of chain elongation
reactions are given below

118

(c) Termination reaction

When the fatty acid residue has attained a desired length the chain elongation
stops at reaction (v) and the cycle is not repeated. The acyl group instead of being
transferred to the SH of the enzyme is transferred to SH group of Co-enzyme A
(CoASH) molecule. Thus, CoA derivate of the fatty acid is produced which can then be
utilized in fat synthesis. The enzyme becomes free.
It is believed that during this process of fatty acid synthesis, the acyl group of fatty acid
bound to the SH group of ACP. The latter then passes it from one enzyme of the
complex to the other.
(3) Condensation of fatty acids and Glycerol
The fats or triglycerides are synthesized not from glycerol and free fatty acids but
from glycerophosphate and CoA derivatives of fatty acid, i.e. fatty acyl CoA residues.
First, there is acylation of glycerophosphate by two fatty acyl CoA molecule to
from phosphatidic acid. Now dephosphorylation occurs in the presence of phosphatase
and a deglyceride is formed. The acylation of the free OH groups of diglyceride
completes the biosynthesis of triglyceride or fat.

119

Lecture No: 20
PHOTOPERIODISM - SHORT DAY, LONG DAY AND DAY NEUTRAL
PLANTS PHYTOCHROME - ROLE OF PHYTOCHROME IN
FLOWERING AND REGULATION OF FLOWERING
PHOTOPERIODISM
Photoperiodism is the phenomenon of physiological changes that occur in plants
in response to relative length of day and night (i.e. photoperiod). The response of the
plants to the photoperiod, expressed in the form of flowering is also called as
photoperiodism. The phenomenon of photoperiodism was first discovered by Garner and
Allard (1920). Depending upon the duration of photoperiod, the plants are classified into
three categories.
1. Short day plants (SDP)
2. Long day plants (LDP)
3. Day neutral plants (DNP)
1. Short day plants

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These plants require a relatively short day light period (usually 8-10 hours) and a
continuous dark period of about 14-16 hours for subsequent flowering. These plants are
also known as long-night plants
E.g. Rice, coffee, soybean, tobacco and chrysanthemum

In short day plants, the dark period is critical and must be continuous. If this dark
period is interrupted with a brief exposure of red light (660-665 nm wavelength),
the short day plant will not flower.

Maximum inhibition of flowering with red light occurs at about the middle of
critical dark period.

However, the inhibitory effect of red light can be overcome by a subsequent

exposure with far-red light (730-735 mm wavelength)

Interruption of the light period with red light does not have inhibitory effect on
flowering in short day plants.

Prolongation of the continuous dark period initiates early flowering.

2. Long day plants

These plants require longer day light period (usually 14-16 hours) in a 24 hours
cycle for subsequent flowering. These plants are also called as short night plants.
E.g. Wheat, radish, cabbage, sugar beet and spinach.

121

In long day plants, light period is critical

A brief exposure of red light in the dark period or the prolongation of light period
stimulates flowering in long day plants.

3. Day neutral plants

These plants flower in all photoperiod ranging from 5 hours to 24 hours
continuous exposure.
E.g. Tomato, cotton, sunflower, cucumber, peas and certain varieties of tobacco.
During recent years, intermediate categories of plants such as long short day
plants and short long day plants have also been recognized.
i. Long short day plants
These are short day plants but must be exposed to long days during early periods
of growth for subsequent flowering. E.g. Bryophyllum.
ii. Short long day plants
These are long day plants but must be exposed to short day during early periods
of growth for subsequent flowering. E.g. certain varieties of wheat and rye.

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Differences between short day and long day plants

1

Short day plant

Plants flower when photoperiod is less

Long day plant

Plants flower when photoperiod is more

than the critical day length

Interruption during light period with

than the critical day length

Interruption during light period with

darkness does not inhibit flowering

Flowering is inhibited if the long dark

darkness inhibit flowering

Flowering occurs if the long dark

period is interrupted by a flash of light

Long continuous and uninterrupted dark

period is interrupted by a flash of light

Dark period is not critical for flowering

period is critical for flowering

Flowering does not occur under

Flowering occurs under alternating

alternating cycles of short day and short

cycles of short day followed by still

light period.

shorter dark periods

Phytochrome
It is observed that that a brief exposure with red light during critical dark period
inhibits flowering in a short day plant and this inhibitory effect can be reversed by a
subsequent exposure with far-red light. Similarly, prolongation of the critical light period
or the interruption of the dark period stimulates flowering in long-day plants.
This inhibition of flowering in short day plant and stimulation of flowering in
long day plants involves the operation of a proteinaceous pigment called phytochrome. It
is present in the plasma membrane of cells and it has two components, chromophore and
protein. Phytochrome is present in roots, coleoptiles, stems, hypocotyls, cotyledons,
petioles, leaf blades, vegetative buds, flower tissues, seeds and developing fruits of higher
plants.
The pigment, phytochrome exists in two different forms i.e., red light absorbing
form which is designated as Pr and far red light absorbing form which is designated as
Pfr. These two forms of the pigment are photo chemically inter convertible. When Pr
form of the pigment absorbs red light (660-665 nm), it is converted into Pfr form. When
Pfr form of the pigment absorbs far red light (730-735 nm), it is converted into Pr form.
The Pfr form of pigment gradually changes into Pr form in dark.
It is considered that during day time, the Pfr form of the pigment is accumulated
in the plants which are inhibitory to flowering in short day plants but is stimulatory in

123

long day plants. During critical dark period in short day plants, this form gradually
changes into Pr form resulting in flowering. A brief exposure with red light will convert
this form again into Pfr form thus inhibiting flowering.
Reversal of the inhibitory effect of red light during critical dark period in SDP by
subsequent far-red light exposure is because, the Pfr form after absorbing far-red light
(730-354 nm) will again be converted back into Pr form.
Prolongation of critical light period or the interruption of the dark period by redlight in long day plants will result in further accumulation of the Pfr form of the pigment,
thus stimulating flowering in long-day plants.
Differences between Pr and Pfr forms of phytochrome
1
2

Pr form
It is blue green in colour
It is an inactive form of phytochrome

Pfr form
It is light green in colour
It is an active form of phytochrome

and it does not show phytochrome

and hence shows phytochrome

mediated responses
mediated responses
It has maximum absorption in red region It has maximum absorption in far-red

(about 680nm)
It can be converted into Pfr form in red

region (about 730nm)

It can be converted into Pr form in far

region (660-665nm)
It is found diffused throughout the

red region (730-735nm)

It is found in discrete areas of cytosol

cytosol
The Pr form contains many double

The Pfr form contains rearranged

bonds in pyrrole rings

double bonds in all pyrrole rings

Significance of photoperiodism
Photoperiodism is an example for physiological preconditioning. The stimulus is
given at one time and the response is observed after months. Exposure to longer
photoperiods hastens flowering (E.g). In wheat, the earing is hastened. During long light
exposure, Pr form is converted into Pfr form and flowering is initiated. If dark period is
greater, Pfr is converted into Pfr form that inhibits flowering.
The important phytochrome mediated photo responses in plants include
photoperiodism, seed germination, sex expression, bud dormancy, rhizome formation,

124

leaf abscission, epinasty, flower induction, protein synthesis, pigment synthesis, auxin
catabolism, respiration and stomatal differentiation.

125

Lecture No: 21
TRANSMISSION OF STIMULUS - THEORIES OF FLOWERING.
Photoperiodic Induction
The influence of the length of day and night on the initiation of flowering is called
photoperiodic induction or photo induction.
Plants may require one or more inductive cycle for flowering. An appropriate
photoperiod in 24 hours cycle constitutes one inductive cycle. If a plant which has
received sufficient inductive cycle is subsequently placed under unfavourable
photoperiod, it will still flower.
Flowering will also occur if a plant receives inductive cycles after intervals of
unfavourable photoperiods (i.e. discontinuous inductive cycle).

This persistence of

photoperiodic after effect is called as photoperiodic induction.

An increase in the number of inductive cycles results in early flowering of the

plant. For instance, xanthium (a short day plant) requires only one inductive
cycle and normally flowers after about 64 days. It can be made to flower even
after 13 days if it has received 4-8 inductive cycle. In such case number of
flowers is also increased.

Continuous inductive cycles promote early flowering than discontinuous

inductive cycle.
Some of the examples of plants which requires more than one inductive cycle for
subsequent flowering are,
Biloxi soybean (SDP) - 2 inductive cycles
Salvia (SDP)

- 17 Inductive cycles

Plantago (LDP)

- 25 Inductive cycles

Critical day length

Maryland mammoth tobacco and xanthium are short day plants, but the Maryland
mammoth tobacco is induced to flower when the photoperiod is shorter than 12 hours

126

(12L /12D) whereas, xanthium is induced to flower when the photoperiod is shorter than
15.5 hours (15.5L /8.5D). The photoperiod required to induce flowering is referred to as
the critical day length. Hence, the critical day length for Maryland mammoth tobacco
and xanthium are 12 and 15.5 hours respectively. A short day plant is one that flowers on
photoperiods shorter than the critical day length.
Long day plants, on the other land, are induced to flower on photoperiods longer
than critical day length. For example, the critical day length for Hyoscyamus niger is 11
hours (11L /13D) and it is induced to flower on photoperiods longer than 11 hours.
Suppose, xanthium and Hyoscyamus niger are exposed to a photoperiod of 14
hours of light and 10 hours of darkness (14L/10D), flowering will be induced in both
plants. Xanthium, a short-day plant, will flower because 14L /10D photoperiod is shorter
than critical day length of 15.5 hours. Hyoscyamus, a long-day plant, will flower because
14L/10D is longer than the critical day length of 11 hours.
Perception of photoperiodic stimulus and presence of a floral hormone

Photoperiodic stimulus is perceived by the leaves and a floral hormone is

produced in the leaves which are then translocated to the apical tip, subsequently
causing initiation of floral primordia.

Photoperiodic stimulus perceived by the leaves can be shown by a simple

experiment on cocklebur (xanthium), a short day plant. Cocklebur plant will
flower if it has previously been kept under short day conditions. If the plant is
defoliated and kept under short day condition, it will not flower. Flowering will
also occur if all the leaves of the plant except one leaf have been removed.

If the cocklebur plant whether intact or defoliated is kept under long day
condition it will not flower. But if even one of its leaves is exposed to short day
condition and the rest are under long day condition, flowering will occur.

The photoperiodic stimulus is transmitted from one branch of the plant to another
branch. For example, if in a two branched cocklebur plant one branch is exposed
to short day and the other to long day photoperiod, flowering occurs on both the
branches.

127

Flowering also occurs if one branch is kept under long day conditions and other
branch from which all the leaves except one have been removed is exposed to
short day condition. However, if one branch is exposed to long photoperiod and
the other has been defoliated, under short day conditions, flowering will not occur
in any of the branches.

Flowering stimulus: Florigen

The flowering stimulus is produced in leaves and translocated to apical and lateral
meristems where flower formation is initiated. Chailakhyan (1937) called the flowering
stimulus or flowering hormone as Florigen.
Flowering stimulus is similar in long day plants and short day plants. This can be
proved by a grafting experiment and can be translocated from one plant to another.
Maryland mammoth tobacco, a short day plant and Hyoscyamus niger, a long day
plant, are grafted so that the leafy shoots of both the species are available for experiment.
If the grafted plants are exposed to either long day a short day conditions, both partners
flower. If grafting union is not formed, the flowering stimulus is not translocated from
one partner to another partner.
Theories of Flowering
1. Bunnings hypothesis
2. Chailakhyans hypothesis
Bunnings hypothesis:
Bunning (1958) assumes the presence of endogenous rhythms (Oscillator which
consist of two half cycles. The first half cycle occurs in day and is called photophilous
phase. During this, anabolic process predominates including flowering in plants. The
other half cycle is dark, sensitive and is called skotophilous phase. In this, catabolic
process (dehydration of starch) predominates.

128

SD plants have a critical day length of 9 hours. This period falls within the
photophilous phase. Light during scotophil phase will inhibit photo process initiated
during photophase. The L.D. plants have a critical day length of 15 hours and some light
falls in the skoto philous phase. Under these conditions in L.D. plants will flower. In
S.D. plants oscillator is present close to skoto philous phase, while in L.D. plants it is
close to photo philous phase.
Chilakhyans hypothesis:
This hypothesis assumes that flowering hormone florigen is a complex of two
types of substances gibberellin and anthesins. Gibberellin is essential for growth of the
plant stems and anthesins are required for flower formation.
According to him, flowering in all annual seed plants requires two phases: (i)
Floral stem formation phase (ii) Flower formation phase. First phase involves increased
carbohydrate metabolism and respiration with increased content of GA in leaves. Second
phase requires intensive nitrogen metabolism, higher content of anthesins in leaves and
nucleic acid metabolites in stem buds.
Long day conditions favour the first phase while short day conditions favour second
phase. In long day plants gibberellins are critical, while anthesins are critical in short day
plants. However, anthesin is hypothetical; it has not been isolated as yet.

129

Lecture No: 22
VERNALISATION - MECHANISM OF VERNALISATION AND ITS
SIGNIFICANCE - DEVERNALISATION

The cold treatment given to plant buds, seeds or seedlings for promoting early
flowering is known as Vernalisation. In short, the chilling treatment for induction of early
flowering is called Vernalisation.
Besides an appropriate photoperiod, certain plants require a low temperature
treatment during their early stages of the life for subsequent flowering in the later stages.
This low temperature treatment requirement was termed vernalization by Lysenko
(1928). Due to vernalization, the vegetative period of the plant is cut short resulting in an
early flowering. In nature, vernalisation takes place in the seed stage in annuals like
winter rye (Secale cereale). The biennials and many perennials respond to cold treatment
at a very late stage. E.g. Henbane, apples etc.
Perception of cold stimulus and presence of floral hormone
The cold stimulus is perceived by the apical meristems. The perception of the
cold stimulus results in the formation of a floral hormone which is transmitted to other
parts of the plant. In certain cases, the cold stimulus may even be transmitted to another
plant across a graft union.
For instance, if a vernalized henbane plant is grafted to an unvernalized henbane
plant, the later also flowers. This is due to the induction of the plant to produce a
hormone named as Vernalin by Melchers (1939).
Conditions necessary for vernalization
1. Age of the plant
The age of the plant is an important factor in determining the responsiveness of
the plant to the cold stimulus and it differs in different species. In cereals like winter

130

wheat, the vernalization is effective only if the germinating seeds have received cold
temperature treatment for sufficient time.
While in the case of biennial variety of henbane (Hyoscyamus niger), the plant
will respond to the cold treatment, only if they are at rosette stage and completed at least
10 days of growth.
2. Appropriate low temperature and duration of the exposure
Most suitable temperature for vernalizing the plants ranges between 1-6C. The
effectiveness of low temperature treatment decreases from 0 to 4C. Low temperature at
about -6C is completely ineffective. Similarly at high temperatures from 7C onwards,
the response of the plants is decreased. Temperature of about 12-14C is almost effective
in vernalizing the plant. Besides an appropriate low temperature, a suitable duration of
the cold treatment is essential for vernalization.

Depending upon the degree of

temperature and in different species this period may vary, but usually the duration of the
chilling treatment is about one and half months or more.
3. Oxygen
The vernalization is an aerobic process and requires metabolic energy. In the
absence of O2, cold treatment becomes completely ineffective.
4. Water
Sufficient amount of water is also essential for vernalization. Vernalization of the
dry seed is not possible.
Mechanism of Vernalization
There are two main theories to explain the mechanism of vernalisation.
1. Phasic developmental theory
This theory was proposed by Lysenko (1934) as follows.
(i)

The growth (increase in size) and development (i.e. progressive change in the
characteristic of the new organs) are two distinct phenomenons.

131

(ii)

According to this theory, the process of the development of an annual seed

plant consists of a series of phases which must occur in some predetermined
sequence.

(iii)

Commencement of any of these phases will take place only when the
preceding phase has been completed.

(iv)

The phases require different external conditions for the completion such as
light and temperature.

(v)

Vernalization accelerates the thermo phase i.e. that phase of development

which is dependent upon temperature.
Thus, in winter wheat, low temperature is required for the completion of first

thermo phase. After this, the next phase that is dependent upon light (photo phase) starts.
Vernalization of winter wheat accelerates the first thermo phase so that there is an early
swing from vegetative to reproductive phase or flowering.
2. Hormonal theories
It has already been described that vernalization probably involves the formation
of a floral hormone called as vernalin. Based on this fact, many hypothetical schemes
have been proposed by different workers from time to time. The first hormonal theory
proposed by Long and Melchers (1947) is schematically shown below.

D
Higher temp.

Cold
A

Normal temp.
B

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Flowering

According to this scheme, the precursor A is converted into a thermo labile

compound B during cold treatment. Under normal conditions B changes into C which
ultimately causes flowering.

But at higher temperature B is converted into D and

flowering does not take place (devernalization).

Devernalization
The positive effect of the low temperature treatment on the vernalization of the
plant can be counteracted by subsequent high temperature. This is called devernalization.
The devernalized plant can again be vernalized by subsequent low temperature treatment.
Vernalization and Gibberellins
The gibberellins are known to replace the low temperature requirement in certain
biennial plants such as henbane, where the plant normally remains vegetative and retains
its rosette habit during the first growing season and after passing through the winter
period flowers in the next season. The gibberellins cause such plants to flower even
during the first year.
Significance of vernalization
1. Vernalization shortens the vegetative period of the plant
2. It increases cold resistance of the plants
3. Vernalization increases the resistance of plants to fungal diseases.
4. It is a physiological process that substitutes or compensates the effect of thermo
phase.
5. In biennials, vernalisation induces early flowering and early fruit setting.
6. A non vernalised shoot apex can be induced to flower by grafting the plant with a
vernalised plant.

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Lecture No: 23
SOURCE SINK RELATIONSHIP - YIELD COMPONENTS - HARVEST
INDEX AND ITS IMPORTANCE
Source
1. It is the regions of photoassimilates production
2. Export photoassimilates
3. Chlorophyllous tissues
4. Leaves, stipules, fruit wall, young stem, pedicel, awns, peduncle, calyx, bract etc
Sink
1. Regions of photoassimilates consumption
2. Import photoassimilates
3. Growing regions
4. Storage organs Fruit and Seed
Source strength
1. Source Size x Source activity
2. Differences in CO2 fixation (Rubisco & PEP Case)
3. Leaf characters size, thickness, mesophyll size, compaction, vascular bundle
4. Carrying capacity of sieve element (temp., H2O, nutrients, hormone)
Sink strength
1. Sink size x Sink activity
2. Potential capacity of the sink to accumulate assimilates
3. Competition among different sink
Source sink interaction
1. Source sink equilibrium
2. Small surplus source for stress
3. High source size during sink differentiation

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4. Improve strength by activity

5. Synchrony of sink organ development
6. Increased HI is reached increase DMA
7. Reduce photorespiration in C3 plants
Evans (1983)
Reduced growth of non harvestable organ
Prolonged faster storage
Enhanced competition of storage organ
Enhanced competition of regulatory process
Reduced stem weight and height
Reduced root weight with adequate nutrient and H2O
Improved agronomic support (avoid biotic & abiotic stress)
Hormonal regulation
Developmental plasticity (small surplus source for stress)
Efficient system
1. Quick export of photoassimilates to avoid end product inhibition
2. Efficient root system
3. More photosynthetic rate
4. Optimum LAI (4 to 6)
5. High photosynthetic rate & high DMA
Blackmans law of limiting factor
1. A process is controlled by several factors
2. The phase of the process is limited by slowest factor
3. Compensation mechanism working under canopy level
DMA
G x E interaction; nutrients; CO2 fixation rate (path way); photorespiration; vascular
network; LAI & LAD; source-sink limiting condition; root-shoot balance

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HI
Ye = Yb x h
/ HI = {Yield (Eco)/ Yield (Biol)} x 100

Improve HI
Increase biomass production (DMA)
Synchronized development of reproductive organ
Reproductively determinate
High source strength at the time of sink differentiation
Reduced growth of non harvestable organ
Reduced leaf growth at reproductive stage with high LAD
Optimum LAI and early peak LAI
More prolonged and faster storage, enhanced competitiveness among of the
storage organ
High photosynthetic rate
Improved HI by increased size and number of sink organ
Decline in duration of Vegetative growth and increased duration of Reproductive
growth.
Limitations
Source: wheat, rice, pulses, oilseeds
Sink: bajra, ragi
Transport: sorghum, maize (green leaf at harvest; senescence of phloem
Parenchyma)
Sink limitation:
Late anthesis (Long Vegetative phase)
Indeterminate (Vegetative & Reproductive growth)
Vegetative growth at Reproductive phase
Less sink number and size
Hormonal imbalance
Any Stress
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Multi-sink demand (nodules supply 25 75 % of N demand)

Source limitation:
Low canopy photosynthesis
Low optimum LAI
Slow peak LAI (lag vegetative growth)
Low LAD at filling
Early leaf senescence
Stress nutrients, water
PGRs:
ABA inhibit sucrose uptake in source (Loading)
Auxin promotes source uptake
Starch accumulation in chloroplast inhibit photosynthesis
ABA in leaves causes closer of stomata (Inhibit CO2 fixation)
Cytokinin delays senescence of source and sink
Cytokinin in sink increases photoassimilates import
Ethylene induces senescence process.

137

Lecture No: 24
GROWTH - GROWTH CURVE, PHASES OF GROWTH AND FACTORS
INFLUENCING GROWTH

Growth is defined as a vital process that brings about a permanent and irreversible
change in any plant or its part in respect to its size, form, weight and volume. Growth is
restricted only to living cells and is accomplished by metabolic processes involving
synthesize of macromolecules, such as nucleic acids, proteins, lipids and polysaccharides
at the expense of metabolic energy.
Growth at cellular level is also accompanied by the organization of
macromolecules into assemblages of membranes, plastids, mitochondria, ribosome and
other cell organelles. Cells do not definitely increase in size but divide, giving rise to
daughter cells. An important process during cell division is synthesis and replication of
nuclear DNA in the chromosomes, which is then passed into the daughter cells.
Therefore, the term growth is used to denote an increase in size by cell division and cell
enlargement, together with the synthesis of new cellulose materials and the organization
of cellular organelles.
Growth regions
Typical growth regions in plants are the apices of shoot and root. Such growing
regions are known as apical meristems, primary meristems or regions of primary growth.
These apical meristems are responsible for the increase in length, differentiation of
various appendages and formation of plant tissues.
Phases of growth
Growth is not a simple process. It occurs in meristematic regions where the
meristematic cell has to pass through the following 3 phases.
1. Cell formation phase
2. Cell elongation phase
3. Cell differentiation (cell maturation)
The cell formation phase is represented by meristematic zone and cell
enlargement phase by cell elongation zone.
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The dividing meristematic cells are thin walled and have dense protoplasm with a
large nucleus and with or without very small vacuoles. The intercellular spaces are also
absent. The newly formed cells after the first phase of cell division have to pass through
the second phase of cell enlargement. During the second phase of cell elongation on
account of large quantities of solutes inside the growing cell, water enters the cell due to
osmotic effect resulting in the increased turgidity and expansion and dilation of the thin
and elastic cell wall. This phase also results in appearance of large vacuoles.
In the last phase or cell maturation, the secondary walls are laid down and cell
matures and gets differentiated into permanent tissue.
Growth curve
Growth curve is a graph obtained by plotting the growth rate of a plant against
time factor.

The growth rate of a cell, a plant organ, a whole plant or the whole life

cycle of plant is measured in terms of length, size, area, volume or weight. It has been
found that different growth phases result in S shaped curve or sigmoid curve. In initial
stages during the phase of cell formation, the growth rate increases slowly while it
increases rapidly during the phase of cell elongation or cell enlargement and again slows
down during the phase of cell maturation.

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The period during which the course of growth takes place is known as grand
period of growth. Thus, in a standard growth curve, three well marked regions can be
observed, the initial growth stage (lag phase), the grand period of growth (exponential or
log phase) and the steady stage (maturity stage or senescence or stationary phase). The
overall growth may be affected by external or internal factors but the S- shaped curve of
grand period of growth is never influenced. This growth curve suits well to the entire life
of an annual plant when measured in terms of dry weight against time.
Early growth of the plant is limited by the amount of food reserves in the seed.
When the emerged seedlings develop an adequate root system and enough leaf surfaces to
support vigorous photosynthesis and anabolism, a period of rapid increase in size is
possible.
High metabolic rates are not maintained indefinitely and eventually processes are
set in motion that leads to cessation of growth. The factors responsible for the decrease
in growth are competition for essential metabolites, growth substances, water, light or the
accumulation of inhibitors, toxic substances or waste materials.
Blackman (1919) suggested that the growth of the plants can be represented by
equation.
W1 = Wo ert
Where, W1 is the final size (Wt, ht etc) after time t. Wo is the initial size at the
beginning of the time period. r is the rate at which plant substance is laid down during
time t and e is the base of natural logarithm. Blackman pointed out that equation
describes the way in which money placed at compound interest increases with time; the
term compound interest law is used to describe such phenomenon. In banks, compound
interest is usually applied quarterly or annually so that the increase in amount occurs as a
jump. With plant system, compound interest is applied continuously and size increase
follows a smooth curve.
From the equation, the final size of an organism (W1) depends on the initial size
(Wo). Larger seed give a larger plant.
In addition, equation shows that plant size also depends on the magnitude of r,
the relative growth rate. Blackman suggested that r might be used as a measure of the

140

ability of the plant to produce new plant material and called r as the efficiency index. The
plants with high efficiency index could be expected to outperform plants with low
efficiency index.
Measurement of growth
The measurement of growth is possible in terms of either increase in weight or
increase in volume or area. The common and simplest method for the measurement of
growth can be a direct method by which the growth is measured by a scale at regular
intervals from beginning to end. The other methods that can be used are horizontal
microscope, auxanometers.
Factors influencing growth
Growth is affected by all factors that affect the activity of protoplasm. Both
physiological and environmental factors such as water, minerals, photosynthesis,
respiration, climate and edaphic factors significantly influence the growth. In general,
factors can be grouped into external and internal factors.
External factors
1. Light
It has direct effect on photosynthesis and transpiration. Light in terms of intensity,
quality and periodicity influence the growth very much.
Light intensity: A weak light promotes shortening of internodes and affects expansion of
leaf. Very weak light reduces the rate of over all growth and also photosynthesis due to
poor development of chlorophyll and higher rate of water loss from the plant.
Light quality: The different wavelengths of light have different responses to growth. In
blue violet radiation, the internodal growth is pronounced while green colour light
promotes the expansion of leaves as compared to complete spectrum of visible light. The
red light favours the growth while infra red and UV is detrimental to growth.
Light duration: The re is remarkable effect of the duration of light on the growth. The
induction and suppression of flowering depend on duration

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2. Temperature
The plants have different temperature requirements based on the region where
they are grown. In general, best growth takes place between 28 and 33 C. and it varies
from temperate to tropical conditions. The optimum temperature requirement is essential
for seed germination, growth, metabolic activities, flowering and yield.
3. Oxygen
The growth of the plant is directly proportional to the amount of oxygen which is
essential for respiration during which the food materials are oxidized to release energy.
4. Carbon dioxide
It is one of the major factors that influence the photosynthesis. The rate of
photosynthesis increases as the availability of CO2 increases while other factors are not
limiting.
5. Water
Water is an essential factor for growth. It is essential for uptake of nutrients,
translocation of nutrients and food materials, regulating transpiration and for various
physiological processes like photosynthesis, respiration and enzymatic activities.
6. Nutrients and food materials
The rate of growth is directly proportional to the availability of nutrients and food
materials. The shortage of food supply affects the growth as it provides the growth
material to the growing region and also it provides the potential energy to the growing
region.
Internal factors
1. Growth hormones and their availability
2. Resistance to climatic, edaphic and biological stresses
3. Photosynthetic rate and respiration
4. Assimilate partitioning and nitrogen content
5. Chlorophyll and other pigments
6. Source-sink relationship and enzyme activities

142

Lecture No: 25
GROWTH ANALYSIS - LAI, LAD, SLW, SLA, LAR, NAR, RGR AND
CGR IN RELATION TO CROP PRODUCTIVITY

Growth analysis
Growth analysis can be used to account for growth in terms that have functional
or structural significance. The type of growth analysis requires measurement of plant
biomass and assimilatory area (leaf area) and methods of computing certain parameters
that describe growth. The growth parameters that are commonly used in agricultural
research and the name of the scientists who proposed the parameters are given below.
LAI

Williams (1946)

LAR

Radford (1967)

LAD -

Power et al. (1967)

SLA

Kvet et al. (1971)

SLW -

Pearce et al. (1968)

NAR -

Williams (1946)

CGR -

Watson (1956)

RGR -

Williams (1946)

HI

Nichiporovich (1951)

i. Leaf Area
This is the area of photosynthetic surface produced by the individual plant over a
period of interval of time and expressed in cm2 plant-1.
ii. Leaf Area Index (LAI)
Williams (1946) proposed the term, Leaf Area Index (LAI). It is the ratio of the
leaf of the crop to the ground area over a period of interval of time. The value of LAI
should be optimum at the maximum ground cover area at which crop canopy receives
maximum solar radiation and hence, the TDMA will be high.

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Total leaf area of a plant

LAI =
Ground area occupied by the plant
iii. Leaf Area Ratio (LAR)
The term, Leaf Area Ratio (LAR) was suggested by Radford (1967), expresses the
ratio between the area of leaf lamina to the total plant biomass or the LAR reflects the
leafiness of a plant or amount of leaf area formed per unit of biomass and expressed in
cm-2 g-1 of plant dry weight.
Leaf area per plant

LAR =
Plant dry weight
iv. Leaf Weight Ratio (LWR)
It was coined by (Kvet et al., 1971) Leaf weight ratio is expressed as the dry
weight of leaves to whole plant dry weight and is expressed in g g 1.
Leaf dry weight

LWR =
Plant dry weight
v. Leaf Area Duration (LAD)
To correlate dry matter yield with LAI, Power et al. (1967) integrated the
LAI with time and called as Leaf Area Duration. LAD takes into account, both the
duration and extent of photosynthetic tissue of the crop canopy. The LAD is expressed in
days.
L1 + L2
LAD =

(t2 t1)

2
L1 = LAI at the first stage
L2 = LAI at the second stage, (t 2 - t 1) = Time interval in days

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vi. Specific Leaf Area (SLA)

Specific leaf area is a measure of the leaf area of the plant to leaf dry weight and
expressed in cm2g-1 as proposed by Kvet et al. (1971).
Leaf area
SLA =
Leaf weight
Hence, if the SLA is high, the photosynthesizing surface will be high. However no
relationship with yield could be expected.
vii. Specific Leaf Weight (SLW)
It is a measure of leaf weight per unit leaf area. Hence, it is a ratio expressed as
g cm-2 and the term was suggested by Pearce et al. (1968). More SLW/unit leaf area
indicates more biomass and a positive relationship with yield can be expected.
Leaf weight
SLW =
Leaf area
viii. Absolute Growth Rate (AGR)
AGR is the function of amount of growing material present and is influenced by
the environment. It gives Absolute values of biomass between two intervals. It is mainly
used for a single plant or single plant organ e.g. Leaf growth, plant weight etc.
h2 h1
cm day-1

AGR =
t2 t1

Where, h1 and h2 are the plant height at t1 and t2 times respectively.

ix. Net Assimilation Rate (NAR)
The term, NAR was used by Williams (1946). NAR is defined as dry matter
increment per unit leaf area or per unit leaf dry weight per unit of time. The NAR is a
measure of the average photosynthetic efficiency of leaves in a crop community.

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(W2 W1)

(loge L2 - loge L1)

NAR =

x
(t2 t1)

(L2 - L1)

Where, W1and W2 is dry weight of whole plant at time t1 and t2 respectively

L1 and L2 are leaf weights or leaf area at t1 and t2 respectively
t1 t2 are time interval in days
NAR is expressed as the grams of dry weight increase per unit dry weight or area per
unit time (g g -1day-1)
x. Relative Growth Rate (RGR)
The term was coined by Williams (1946). Relative Growth Rate (RGR) expresses
the total plant dry weight increase in a time interval in relation to the initial weight or Dry
matter increment per unit biomass per unit time or grams of dry weight increase per gram
of dry weight and expressed as unit dry weight / unit dry weight / unit time (g g -1day-1)
loge W2 loge W1
RGR =
t2 t1
Where, W1 and W2 are whole plant dry weight at t1 and t2 respectively
t1 and t2 are time interval in days

xi. Crop Growth Rate (CGR)

The method was suggested by Watson (1956). The CGR explains the dry
matter accumulated per unit land area per unit time (g m-2 day-1)
(W2 W1)
CGR =
(t2 t1)
Where, W1 and W2 are whole plant dry weight at time t1 t2 respectively
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 is the ground area on which W1 and W2 are recorded.

CGR of a species are usually closely related to interception of solar radiation
xii. Total dry matter production (TDMP) and its distribution
The TDMP is the biomass accumulated by the whole plant over a period of
interval of time and its distribution (allocation) to different parts of the plant such as
roots, stems, leaves and the economic parts which controls the sink potential.
xiii. Translocation percentage (TP)
The term translocation percentage indicates the quantum of photosynthates
translocated from source (straw) to the grain (panicle/grains) from flowering to harvest.
Straw weight at flowering straw weight at harvest
TP =
Panicle weight at flowering panicle weight at harvest
xiv. Light extinction coefficient
It is the ratio of light intercepted by crop between the top and bottom of crop
canopy to the LAI.
loge I / Io
K=
LAI
Where, Io and I are the light intensity at top and bottom of a population with LAI
xv. Light Transmission Ratio (LTR)
It is expressed as the ratio of quantum of light intercepted by crop canopy at top to
the bottom. Light intensity is expressed in K lux or W m-2
LTR = I / Io
Where, I : light intercepted at the bottom of the crop canopy
Io: light intercepted at the top of the crop canopy

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xvi. Dry Matter Efficiency (DME)

It is defined as the percent of dry matter accumulated in the grain from the total
dry matter produced over the crop growth period.
Grain yield
DME =

100
x

TDMP

Duration of crop

xvii. Unit area efficiency (UAE)

It is expressed as the quantum of grain yield produced over a unit land area for a
specified crop growth period.
Grain yield
UAE =

1
x

Land area

Duration of crop

xviii) Harvest Index

The harvest index is expressed as the percent ratio between the economic yield
and total biological yield and was suggested by Nichiporovich (1951).
Economic yield
HI =

x 100
Total biological yield

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Lecture No: 26
PLANT GROWTH REGULATORS - GROWTH HORMONES DEFINITION AND CLASSIFICATION - PHYSIOLOGICAL ROLE
OF AUXINS AND GA.

PLANT GROWTH REGULATORS

Plant growth regulators or phytohormones are organic substances produced
naturally in higher plants, controlling growth or other physiological functions at a site
remote from its place of production and active in minute amounts. Thimmann (1948)
proposed the term Phyto hormone as these hormones are synthesized in plants. Plant
growth regulators include auxins, gibberellins, cytokinins, ethylene, growth retardants
and growth inhibitors. Auxins are the hormones first discovered in plants and later
gibberellins and cytokinins were also discovered.

Hormone
An endogenous compound, which is synthesized at one site and transported to
another site where it exerts a physiological effect in very low concentration. But ethylene
(gaseous nature), exert a physiological effect only at a near a site where it is synthesized.
Classified definition of a hormone does not apply to ethylene.
Plant growth regulators

Defined as organic compounds other than nutrients, that affects the

physiological processes of growth and development in plants when applied in
low concentrations.

Defined as either natural or synthetic compounds that are applied directly to a

target plant to alter its life processes or its structure to improve quality,
increase yields, or facilitate harvesting.

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Plant Hormone
When correctly used, is restricted to naturally occurring plant substances, there
fall into five classes. Auxin, Gibberellins, Cytokinin, ABA and ethylene. Plant growth
regulator includes synthetic compounds as well as naturally occurring hormones.
Plant Growth Hormone
The primary site of action of plant growth hormones at the molecular level
remains unresolved.
Reasons

Each hormone produces a great variety of physiological responses.

Several of these responses to different hormones frequently are similar.

The response of a plant or a plant part to plant growth regulators may vary with the
variety of the plant.

Even a single variety may respond differently depending on its age, environmental
conditions and physiological state of development (especially its natural hormone
content) and state of nutrition.

There are always exceptions for a general rule

suggesting the action of a specific growth regulator on plants.

There are several proposed modes of action in each class of plant hormone, with
substantial arguments for and against each mode.

Hormone groups
Auxin

Substances generally resembles IAA and has the ability

to stimulate the elongation of coleoptiles.

Gibberellins

are diterpenoids, which have the ability to elongate the

stem of green seedlings especially certain dwarf and rosette
types.

Cytokinin

Usually substituted Adenines, which resembles zeatin

(Naturally occurring cytokinin in Zea mays) and have the
ability to stimulate cytokinensis in cultures of tobacco cells.

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Ethylene

Gaseous regulator that stimulate is diametric growth

in the apices of dicot seedlings.

Inhibitors

are regulators of growth, which originally depress the

Auxins
Auxins are a group of phytohormones produced in the shoot and root apices and
they migrate from the apex to the zone of elongation. Auxins promote the growth along
the longitudinal axis of the plant and hence the name (auxeing : to grow). The term, auxin
was introduced by Kogl and Haagen- Smit (1931). Went (1928) isolated auxin from the
Avena coleoptile tips by a method called Avena coleoptile or curvature test and
concluded that no growth can occur without auxin. Auxins are widely distributed through
out the plant however, abundant in the growing tips such as coleoptile tip, buds, root tips
and leaves. Indole Acetic Acid (IAA) is the only naturally occurring auxin in plants. The
synthetic auxins include,

Avena Curvature Test

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IBA

: Indole Butyric Acid

NAA : Naphthalene Acetic acid

MENA: Methyl ester of Naphthalene acetic acid
MCPA: 2 Methyl 4 chloro phenoxy acetic acid
TIBA : 2, 3, 5 Tri iodo benzoic acid
2, 4-D : 2, 4 dichloro phenoxy acetic acid
2, 4, 5-T: 2, 4, 5 Trichloro phenoxy acetic acid
Natural auxins may occur in the form of either free auxins- which freely move or
diffuse out of the plant tissues readily or bound auxins- which are released from plant
tissues only after hydrolysis, autolysis or enzymolysis.
Physiological effects of auxin
1. Cell division and elongation
The primary physiological effects of auxin are cell division and cell elongation in
the shoots. It is important in the secondary growth of stem and differentiation of xylem
and phloem tissues.
2. Apical dominance
In many plants, if the terminal bud is intact and growing, the growth of lateral
buds just below it remains suppressed. Removal of the apical bud results in the rapid
growth of lateral buds. This phenomenon in which the apical bud dominates over the
lateral buds and does not allow the lateral buds to grow is known as apical dominance.
Skoog and Thimmann (1948) pointed out that the apical dominance might be
under the control of auxin produced at the terminal bud and which is transported
downward through the stem to the lateral buds and hinders the growth. They removed
the apical bud and replaced it with agar block. This resulted in rapid growth of lateral
buds. But when they replaced the apical bud with agar block containing auxin, the lateral
buds remained suppressed and did not grow.

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3. Root initiation
In contrast to stem, the higher concentration of auxin inhibits the elongation of
roots but the number of lateral roots is considerably increased i.e., higher concentration of
auxin induces more lateral branch roots. Application of IAA in lanolin paste (lanolin is a
soft fat prepared from wool and is good solvent for auxin) to the cut end of a young stem
results in an early and extensive rooting. This fact is of great practical importance and
has been widely utilized to promote root formation in economically useful plants which
are propagated by cuttings.
4. Prevention of abscission
Natural auxins prevent the formation of abscission layer which may otherwise
result in the fall of leaves, flowers and fruits.
5. Parthenocarpy
Auxin can induce the formation of parthenocarpic fruits (fruit formation without
pollination and fertilization). In parthenocarpic fruits, the concentration of auxin in the
ovaries is higher than in the ovaries of plants which produce fruits only after fertilization.

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In the later cases, the concentration of the auxin in ovaries increases after pollination and
fertilization.
6. Respiration
Auxin stimulates respiration and there is a correlation between auxin induced
growth and respiration. Auxin may increase the rate of respiration indirectly through
increased supply of ADP by rapidly utilizing ATP in the expanding cells.
7. Callus formation
Besides cell elongation, auxin may also be active in cell division. In many tissue
cultures, where the callus growth is quite normal, the continued growth of such callus
takes place only after the addition of auxin.
8. Eradication of weeds
Some synthetic auxins especially 2, 4- D and 2, 4, 5-T are useful in eradication of
weeds at higher concentrations.
9. Flowering and sex expression
Auxins generally inhibit flowering but in pine apple and lettuce it promotes
uniform flowering.
Distribution of auxin in plants
In plants, auxin (IAA) is synthesized in growing tips or meristematic regions from
where; it is transported to other plant parts. Hence, the highest concentration of IAA is
found in growing shoot tips, young leaves and developing auxiliary shoots. In monocot
seedling, the highest concentration of auxin is found in coleoptile tip which decreases
progressively towards its base.
In dicot seedlings, the highest concentration is found in growing regions of shoot,
young leaves and developing auxiliary shoots. Within the plants, auxin may present in
two forms. i.e., free auxins and bound auxins. Free auxins are those which are easily
extracted by various organic solvents such as diethyl ether. Bound auxins on the other
hand, need more drastic methods such as hydrolysis, autolysis, enzymolysis etc. for

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extraction of auxin. Bound auxins occur in plants as complexes with carbohydrates such
as glucose, arabionse or sugar alcohols or proteins or amino acids such as aspartate,
glutamate or with inositol.
Biosynthesis of auxin (IAA) in plants
Thimann (1935) found that an amino acid, tryptophan is converted into Indole 3
acetic acid. Tryptophan is the primary precursor of IAA in plants. IAA can be formed
from tryptophan by two different pathways.
1. By deamination of tryptophan to form indole-3-pyruvic acid followed by
decarboxylation to from indole-3-acetaldehyde.

The enzymes involved are

tryptophan deamintion and indole pyruvate decarboxylase respectively.

2. By decarboxylation of tryptophan to form tryptamine followed by deamination to
form indole-3-acetaldehyde

and the enzymes

involved are tryptophan

decarboxylase and tryptamine oxidase respectively. Indole

3-acetaldehyde

can

readily be oxidized to indole 3-acetic acid (IAA) in the presence of indole 3acetaldehyde dehydrogenase.
Transport of auxin in plant
The transport of auxin is predominantly polar. In stems, polar transport of auxin
is basipetal i.e., it takes place from apex towards base. Polar transport of auxin is
inhibited by 2, 3, 5 Triiodobenzoic acid (TIBA) and Naphthyl thalamic acid (NPA). The
substances are called as antiauxins.
Destruction / Inactivation of auxin in plants
Auxin is destructed by the enzyme IAA oxidase in the presence of O 2 by
oxidation.
IAA Oxidase
IAA + H2O2 + O2

3-methylene oxindole + H2O + CO2

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Rapid inactivation may also occur by irradiation with x-rays and gamma rays.
UV light also reduces auxin levels in plants. Inactivation or decomposition of IAA by
light has been called as photo oxidation.
Mechanism of Action
IAA increases the plasticity of cell walls so that the cells stretch easily in response
to turgor pressure. It has been suggested that IAA acts upon DNA to influence the
production of mRNA. The mRNA codes for specific enzymes responsible for expansion
of cell walls. Recent evidences indicate that IAA increases oxidative phosphorylation in
respiration and enhanced oxygen uptake. The growth stimulation might be due to
increased energy supply and it is also demonstrated that auxin induces production of
ethylene in plants.
Gibberellins
Discovery
A Japanese scientist Kurosawa found that the rice seedlings infected by the fungus
Gibberella fujikuroi grow taller and turned very thin and pale. An active substance was
isolated from the infected seedlings and named as Gibberellin.
Biosynthesis of gibberellins in plants
The primary precursor for the formation of gibberellins is acetate.
Acetate + COA Acetyl COA Mevalonic acid MA pyrophosphate
Isopentanyl pyrophosphate Geranyl pyrophosphate GGPP Kaurene
Gibberellins.
Physiological effects of gibberellins
1. Seed germination
Certain light sensitive seeds eg. Lettuce and tobacco show poor germination in
dark. Germination starts vigorously if these seeds are exposed to light or red light. This
requirement of light is overcome if the seeds are treated with gibberellic acid in dark.

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2. Dormancy of buds
In temperature regions the buds formed in autumn remain dormant until next
spring due to severe cold.
treatments.

This dormancy of buds can be broken by gibberellin

In potato also, there is a dormant period after harvest, but the application

of gibberellin sprouts the refer vigorously.

3. Root growth
Gibberellins have little or no effect on root growth. At higher concentration, some
inhibition of root growth may occur. The initiation of roots is markedly inhibited by
gibberellins in isolated cuttings.

4. Elongation of internodes
The most pronounced effect of gibberellins on the plant growth is the elongation
of the internodes.

Therefore in many plants such as dwarf pea, dwarf maize etc

gibberellins overcome the genetic dwarfism.

5. Bolting and flowering

In many herbaceous plants, the early period of growth shows rosette habit with
short stem and small leaves. Under short days, the rosette habit is retained while under
long days bolting occurs i.e. the stem elongates rapidly and is converted into polar axis
bearing flower primordia.

This bolting can also be induced in such plants by the

application of gibberellins even under non-inductive short days.

In Hyoscyamus niger (a long day plant) gibberellin treatment causes bolting and
flowering under non-inductive short days. While in long day plants the gibberellin
treatment usually results in early flowering. In short day plants, its effects are quite
variable. It may either have no effect or inhibit or may activate flowering.

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6. Parthenocarpy
Germination of the pollen grains is stimulated by gibberellins; likewise, the
growth of the fruit and the formation of parthenocarpic fruits can be induced by
gibberellin treatment. In many cases, eg. pome and stone fruits where auxins have failed
to induce parthenocarpy, the gibberellins have proven to be successful. Seedless and
fleshly tomatoes and large sized seedless grapes are produced by gibberellin treatments
on commercial scale.

7. Synthesis of the enzyme - amylase

One important function of gibberellins is to cause the synthesis of the enzyme amylase in the aleurone layer of the endosperm of cereal grains during germination. This
enzyme brings about hydrolysis of starch to form simple sugars which are then
translocated to growing embryo to provide energy source.

Distribution of gibberellins in plant

Gibberellins are found in all parts of higher plants including shoots, roots, leaves,
flower, petals, anthers and seeds. In general, reproductive parts contain much higher
concentrations of gibberellins than the negative parts. Immature seeds are especially rich
in gibberellins (10-100 mg per g fresh weight).
In plants, gibberellins occur in two forms free gibberellins and bound gibberellins.
Bound gibberellins usually occur as gibberellin glycosides.

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Lecture No: 27
PHYSIOLOGICAL ROLE OF CYTOKININ, ETHYLENE AND ABA SYNTHETIC GROWTH REGULATORS AND THEIR USES IN
CROP PRODUCTIVITY.
CYTOKININS (Kinetin)
Kinetin was discovered by Skoog and Miller (1950) from the tobacco pith callus
and the chemical substance was identified as 6-furfuryl aminopurine. Because of its
specific effect on cytokinesis (cell division), it was called as cytokinins or kinetin. The
term, cytokinin was proposed by Letham (1963). Fairley and Kingour (1966) used the
term, phytokinins for cytokinins because of their plant origin. Chemically cytokinins are
kinins and they are purine derivatives.
Cytokinins, besides their main effect on cell division, also regulate growth and
hence they are considered as natural plant growth hormones. Some of the very important
and commonly known naturally occurring cytokinins are Coconut milk factor and Zeatin.
It was also identified that cytokinin as a constituent of t-RNA.
Naturally occurring cytokinins
Cytokinins can be extracted from coconut milk (liquid endosperm of coconut),
tomato juice, flowers and fruits of Pyrus malus; fruits of Pyrus communis (Pear), Prunus
cerasiferae (plum) and Lycopersicum esculentum (bhendi); Cambial tissues of Pinus
radiata, Eucalyptus regnans and Nicotiana tabacum; immature fruits of Zea mays,
Juglans sp. and Musa sp; female gametophytes of Ginkgo biloba; fruitlets, embryo and
endosperms of Prunus persica; seedling of Pisum sativum; root exudates of Helianthus
annuus and tumour tissues of tobacco. According to Skoog and Armstrong (1970), at
least seven well established types of cytokinins have been reported from the plants.
Biosynthesis
It is assumed that cytokinins are synthesised as in the case of purines in plants
(nucleic acid synthesis). Root tip is an important site of its synthesis. However,

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developing seeds and cambial tissues are also the site of cytokinin biosynthesis. Kende
(1965) reported that cytokinins move upwards perhaps in the xylem stream. However,
basipetal movement in petiole and isolated stems are also observed. Seth et al (1966)
found that auxin enhances kinetin movement (translocation) in bean stems.
Physiological effects of cytokinins
1. Cell division
The most important biological effect of kinetin on plants is to induce cell division
especially in tobacco pith callus, carrot root tissue, soybean cotyledon, pea callus etc.
2. Cell enlargement
Like auxins and gibberellins, the kinetin may also induce cell enlargement.
Significant cell enlargement has been observed in the leaves of Phaseolus vulgaris,
pumpkin cotyledons, tobacco pith culture, cortical cells of tobacco roots etc.
3. Concentration of apical dominance
External application of cytokinin promotes the growth of lateral buds and hence
counteracts the effect of apical dominance
4. Dormancy of seeds
Like gibberellins, the dormancy of certain light sensitive seeds such as lettuce and
tobacco can also be broken by kinetin treatment.
5. Delay of senescence ( Richmand - Lang effect)
The senescence of leaves usually accompanies with loss of chlorophyll and rapid
breakdown of proteins. Senescence can be postponed to several days by kinetin treatment
by improving RNA synthesis followed by protein synthesis.
Richmand and Lang (1957) while working on detached leaves of Xanthium found
that kinetin was able to postpone the senescence for a number of days.
6. Flower induction
Cytokinins can be employed successfully to induce flowering in short day plants.

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7. Morphogenesis
It has been shown that high auxin and low kinetin produced only roots whereas
high kinetin and low auxin could promote formation of shoot buds.
8. Accumulation and translocation of solutes
Plants accumulate solutes very actively with the help of Cytokinin and also help
in solute translocation in phloem.
9. Protein synthesis
Osborne (1962) demonstrated the increased rate of protein synthesis due to
translocation bys kinetin treatment.
10. Other effects
Cytokinins provide resistance to high temperature, cold and diseases in some
plants. They also help in flowering by substituting the photoperiodic requirements. In
some cases, they stimulate synthesis of several enzymes involved in photosynthesis.
11. Commercial applications
Cytokinins have been used for increasing shelf life of fruits, quickening of root
induction and producing efficient root system, increasing yield and oil contents of oil
seeds like ground nut.
Ethylene
Ethylene is the only natural plant growth hormone exists in gaseous form.

Important physiological elects

1. The main role of ethylene is it hastens the ripening of fleshy fruits eg. Banana,
apples, pears, tomatoes, citrus etc.
2. It stimulates senescence and abscission of leaves
3. It is effective in inducing flowering in pine apple

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4. It causes inhibition of root growth

5. It stimulates the formation of adventitious roots
6. It stimulates fading of flowers
7. It stimulates epinasty of leaves.

Abscisic acid
Addicott (1963) isolated a substance strongly antagonistic to growth from young
cotton fruits and named Abscissin II. Later on this name was changed to Abscisic acid.
This substance also induces dormancy of buds therefore it also named as Dormin.
Abscisic acid is a naturally occurring growth inhibitor.

Physiological effects
The two main physiological effects are
1. Geotropism in roots
2. Stomatal closing
Besides other effects

1. Geotropism in roots
Geotropic curvature of root is mainly due to translocation of ABA in basipetal
direction towards the root tip.

2. Stomatal closing
ABA is synthesized and stored in mesophyll chloroplast. In respond to water
stress, the permeability of chloroplast membrane is lost which resulted is diffusion of
ABA out of chloroplast into the cytoplasm of the mesophyll cells. From mesophyll cells
it diffuses into guard cells where it causes closing of stomata.

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3. Other effects
i. Including bud dormancy and seed dormancy
ii. Includes tuberisation
iii. Induces senescence of leaves fruit ripening, abscission of leaves, flowers and fruits
iv. Increasing the resistance of temperate zone plants to frost injury.

Growth retardants
There is no. of synthesis compounds which prevent the gibberellins from
exhibiting their usual responses in plants such as cell enlargement or stem elongation. So
they are called as anti gibberellins or growth retardants. They are
1. Cycocel (2- chloroethyl trimethyl ammonium chloride (CCC)
2. Phosphon D (2, 4 dichlorobenzyl tributyl phosphonium chloride)
3. AMO 1618
4. Morphactins
5. Maleic hydrazide

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Lecture No: 28
PRACTICAL APPLICATION OF PLANT GROWTH REGULATORS IN
CROP PRODUCTIVITY
Commercial uses of growth regulators
1. Rooting and plant propagation
a) Auxin compound like IBA NAA, 2,4-D, 2, 4,5-T
b) IBA produces strong fibrous root system
2. Germination and dormancy
a) Gibberellin is a potent germination promoter
b) Abscissic acid germination inhibitor (Anti Gibberellin)
c) Induce Dormancy

- ABA

d) Breaking of dormancy - Auxins and Gibberellin

3. Fruit set and Development
a) Fruit setting

2, 4, 5 T

b) Fruit size increment in grapes

c) Shelf life increment in fruits and flowers

Gibberellic acid

Cytokinin

d) Good fruit shape ------ Gibberellic acid + Cytokinin

e) Parthenocarpic fruit Gibberellins, IAA and PAA
4. Sex expression
Production of male flowers

Gibberellins (cucumber)

Production of female flowers

Auxins and Gibberellins

Cucumber
maize

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5. Abiscission
Control of abscission

NAA and IAA

Induce Abscission

Ethrel

6. Morphogenesis
Auxin and Cytokinin
7. Weed control
2, 4-D and 2, 4, 5-T
8. Plant organ size
Increases plant height

GA

Shorten the plant height

TIBA

Tri iodo benzoic acid

Increases Tillering

Cytokinin
Ex: BAP (Benzyl amino purine) and TIBA

9. Antitranspirants

ABA and PMA

Phenyl mercury acetate

10. Papaya Later flow

Ethephon

11. Rubber latex flow

2, 4 D and 2, 4,5 T

12. Fruit ripening

Ethrel

13. Sugarcane ripeners

Glyphosphate and CCC

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166

Lecture No: 29

Environmental stresses - water stress - physiological

changes - adaptation to drought and amelioration.

I. DROUGHT (Water stress)

Drought is defined as the deficiency of water severe enough to check the plant
growth. Drought has been classified into two broad categories viz., soil drought and
atmospheric drought. Soil drought leads to atmospheric drought. Atmospheric drought
occurs due to low atmospheric humidity, high wind velocity and high temperature which
cause a plant to lose most of its water.
Physiological changes occur due to drought
1. Functioning of stomata
In general, stomata lose their function and may die, because wilting after certain
limit denatures the starch in the guard cells and also in the mesophyll cells.

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2. Carbohydrates metabolism in green leaves

The very first effect of drought on carbohydrates metabolism is that starch
disappears from the wilted leaves and sugar accumulates simultaneously.
3. Photosynthetic activity
CO2 diffusion into the leaf is prevented due to decrease in stomatal opening and
there by reduces photosynthetic activity in green cells.
4. Osmotic pressure
The reduced amount of water during drought causes an increase in the osmotic
pressure of plant cell. This increase in osmotic pressure permits the plant to utilize better
soil moisture.
5. Permeability
The permeability to water and urea increases during drought.
6. Biochemical effects
Water shortage alters the chemical composition. For example, starch is converted
to sugar, besides this, there is a considerable increase in nitrate nitrogen and protein
synthesis is adversely affected.
Adaptation to drought
Drought resistance
Drought resistance is defined as the capacity of plants to survive during the
period of drought with little or no injury. There are three important categories of plants
growing in the areas facing drought. They are ephemerals, succulents and non-succulent
perennials
1. Ephemerals
These are short lived plants and they complete their life cycle within a short
favourable period during rainy season. They pass dry periods in the form of seeds. They
are called as drought escaping plants.
2. Succulent plants
These plants accumulate large quantities of water and use it slowly during dry
period. Thus, they pass dry periods or drought without facing it. Such plants develop

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several morphological adaptations for reducing transpiration such as thick cuticle,

reduced leaf area, sunken stomata etc.
3. Non succulent plants
These plants are in fact the real drought enduring (tolerant) plants. They tolerate
drought without adapting any mechanism to ensure continuous supply of water. They
develop many morphological adaptations which are collectively called xeromorphy. They
develop, in general, greyish colour, reflecting surfaces, smaller leaves, extensive root
system, leaf fall during dry season, sunken stomata and thick cuticle etc. They develop
an elaborated conducting system. The stomata remain closed mostly in dry periods.
The plants develop several protoplasmic peculiarities such as cell size, cell
structure, increased permeability, increased imbibition power, elasticity, small vacuoles,
higher osmotic pressure etc.

Osmotic adjustment

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Methods to overcome drought

Selection of drought tolerant species

Adjusting the tome of sowing in such a way that the crop completes its lifecycle
before the onset of drought

Seed hardening with KCl, KH2PO4, CaCl2 or Thiourea

Thinning of poorly established plants

Mulching to minimize the evaporative loss

Foliar spray of antitranspirants such as Kaolin, PMA, Waxes and Silicone oils

Foliar spray of KCl

Foliar spray of growth retardants such as CCC and MC

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Lecture No: 30

Temperature stress - Physiological changes - low and high

temperature - chilling injury - tolerance - alleviation.
II. TEMPERATURE STRESS
Temperature stress includes both high temperature stress and low temperature
stress. Low temperature stress causes chilling injury and freezing injury.
1. Chilling injury
The tropical origin plants are injured when the temperature drops to some point
close to 0C. The injury which occurs due to low temperature but above zero degree
centigrade is called chilling injury.
2. Freezing injury
Freezing injury occurs when the temperature is 0C or below.
Effect of freezing and chilling injury plants

The lipid molecules in cell membrane get solidified i.e. changed from liquid state
to solid state. Hence, the semi-permeable nature of the membrane is changed and
the membrane becomes leaky.

Inactivation of mitochondria

Streaming of protoplasm is stopped

Accumulation of respiratory metabolites which become highly toxic

Ice formation inside the cell occurs.

Prevention of cold injury

Some plants change the pattern of growth.

The growth is completely arrested during this period.

In cell membrane, unsaturated fatty acid content is increased.

171

Intracellular ice formation is reduced.

The quantity of free enzymes, sugars and proteins increases.

High temperature stress

The effect of high temperature is heat Injury. Heat Injury occurs when plant
temperature is higher than that of environment (exceeds 35C).
General effects of high temperature
High temperature affects
1. Seedling growth and vigour
2. Water and nutrient uptake
3. Solute transport
4. Photosynthesis and respiration
5. General metabolic processes
6. Fertilization and maturation

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Lecture No: 31

Low light and UV radiation stresses - salt stress physiological changes and alleviation.
IV. LOW LIGHT STRESS
In some places (e.g. Thanjavur), the light intensity might be even up to 60000 lux
in the first season but it would be low up to 30000 lux in the second season causing very
poor productivity. Light quality is also very poor by showing about 400-440nm instead of
the normal 600-640nm. The abnormal light intensity and quality causes reduced yield in
any crops.
V. UV-RADIATION STRESS
UV radiation is divided into three categories
1. UV A wavelength ranges from 320 to 400 nm and this is less lethal to the plants.
2. UV B wavelength ranges from 280 to 320 nm and this is lethal to the plants.
3. UV C wavelength is less than 280 nm and it is highly lethal to all biological
systems.
The UV radiations cause environmental stress as the cell constituents like proteins
and nucleic acids absorb UV radiation in the range of 250-400 nm (UV A and UV B) and
cause death of the tissues. In general, on the outer atmosphere of the earth, CO2, ozone
and water vapour form a layer and this layer prevent the entry of UV radiation. However,
ozone depletion causes easy entry of UV radiation. In general, 1% reduction in ozone
(O3) causes 2% increase in UV radiation.
UV radiation and plant response
1. UV radiation slows down the growth of plants
2. Damage the process of photosynthesis
3. Prevent maturation and ripening process
4. Accelerate genetic mutation.
III. SALT STRESS
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Salt stress occurs due to excess salt accumulation in the soil. As a result, water
potential of soil solution decreases and therefore exosmosis occurs.

This leads to

physiological drought causing wilting of plants.

Classification of saline soil: 1. Saline soil 2. Alkaline soil
1. Saline soil
In saline soils, the electrical conductivity is greater than 4 dS/m, exchangeable
sodium percentage is less than 15% and pH is less than 8.5. These soils are dominated by
Cl- and SO2-4 ions.
2. Alkaline soil
Alkaline soils are also termed as sodic soils wherein, the electrical conductivity is
less than 4 dS/m, exchangeable sodium percentage is greater than 15% and pH of the soil
is greater than 8.5. These soils are dominated by CO-3 and HCO-3 ions.
Classification of plants
Plants are classified into two types based on the tolerance to salt stress. They are
halophytes and glycophytes.
1. Halophytes
Halophytes are the plants that grow under high salt concentrations. They are again
divided into two types based on extreme of tolerance.
Euhalophytes: can tolerate extreme salt stress
Oligohalophytes: can tolerate moderate salt stress
2. Glycophytes
Glycophytes are the plants that cannot grow under high salt concentration.

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Effect of salt stress on plant growth and yield

1. Seed germination
Salt stress delays seed germination due to the reduced activity of the enzyme, amylase
2. Seedling growth
The early seedling growth is more sensitive. There is a significant reduction in
root emergence, root growth and root length.
3. Vegetative growth
When plants attain vegetative stage, salt injury is more severe only at high
temperature and low humidity. Because under these conditions, the transpiration rate will
be very high as a result uptake of salt is also high.
4. Reproductive stage
Salinity affects panicle initiation, spikelet formation, fertilization and pollen grain
germination.
5. Photosynthesis
Salinity drastically declines photosynthetic process. Thylakoid are damaged by
high concentration of salt and chlorophyll b content is drastically reduced.
Mechanism of salt tolerance
1. Some plants are able to maintain high water potential by reducing the
transpiration rate.
2. Salts are accumulated in stem and older leaves in which metabolic processes take
place in a slower rate.
3. Na+ (sodium ion) toxicity is avoided by accumulating high amount of K+ ions.
4. Accumulation of toxic ions in the vacuole but not in the cytoplasm.
5. Accumulation of proline and abscissic acid which are associated with tolerance of
the plants to salt.
Relative salt tolerant crops
Tolerant crops: Cotton, sugar cane, barley
Semi tolerant crops: Rice, maize, wheat, oats, sunflower, soybean
Sensitive crops: Cow pea, beans, groundnut and grams

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Management of salt stress

1. Leaching of salts with adequate water
2. Application gypsum to convert the highly injurious carbonates to less injurious
sulphate
3. Selection of salt tolerant crops
4. Use of FYM and other organic manures

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Lecture No: 32

Global warming - physiological effects on crop

productivity
VI. GLOBAL WARMING (Green house Effect)
In general, delicate plants which require protection from weather are grown in
green house (glass house). In green house so many gases are produced like CO 2, water
vapour, methane, oxides of nitrogen and chloro fluoro carbon (CFC). These gases are
produced from plants and accumulated inside the glass house; as a result glass house gets
warming. In natural atmosphere also the same effect occurs i.e. global warming (due to
the release of gases from plants).
But in glass house, glass roof is present to prevent the escape of gases from the
glass house. In natural atmosphere, the gases such as ozone, water vapour, CO 2 methane
etc. form a layer on the lower atmosphere and this layer prevents the heat escaping from
the earth. If heat is released or escaped from earth, the temperature of earth would be
below freezing point. The accumulation of heat or gases causes the warming of earth
surface and leads to global warming.
Global warming leads to the following effects:
1.

Rise in temperature

2.

Average rise in the level of sea (about 6 cm/decade) due to melting of polar ice.

3.

Steady increase (enrichment) in the CO2.

Atmosphere (5.1 x 1018kg)

Lithosphere - (1.5 x 1022 kg)

are in a dynamic equilibrium

Hydrosphere 1.4 x 10 22 kg)

Biosphere 1.2 x 1015 kg (dry wt.)
CO2 fixation by green plants
Total area of green plants 510 x106 km2
CO2 fixed 1.39 x 1014 kg year-1 (0.27 kg m-2 year-1)
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Light utilization:
A small portion of radiant energy (400 700 nm) is reaching the earths surface
0.1 1.0 % utilized under natural vegetation or ordinary agriculture
2.0 2.5 % under intensive agriculture
6.0 10.0 % in some crop plants
20.0 25.0 % under laboratory condition
Biosphere:
O2 in atmosphere

- 1.1 x 1018 kg

O2 released by photosynthesis- 5.1 x 1014 kg year-1

CO2 content

- 5 x 1016 kg (mostly dissolved in sea)

CO2 consumed by photosynthesis

- 7 x 1014 kg year-1

Human activities disturbed the global ecosystem (MST- mesospheric,

Stratospheric, Troposphere)
Landscape modification
Resource exploitation
Effluent flow
High temperature
Rainfall redistribution
Increased UV-B radiation due to stratospheric O3 depletion
Increased level of atmospheric CO2
Other green house gases
Transparent to incoming short wave radiation
Absorb short wave and emit long wave radiation
Net emission of CO2
Bacterial fermentation in the anaerobic rice fields generate 120 million ton CH4
every year
Ruminant gut bacteria produce 78 m tons of CH4 every year and are released by
Flatulence
Green house gases
CO2, CH4, NO, NO2, N2O, (NxOx), CFCl3, CF2Cl2, CFMs, O3, H2O

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Fossil fuel reserves are large enough for climatic changes to occur, if these
reserves continue to be exploited at a higher rate in future

CO2 enrichment and crop productivity

1. CO2 enrichment leads to increased photosynthesis and productivity
2. CO2 enrichment also decreases stomatal conductance by closing the stomata, thee
by decrease the transpiration / unit area of the leaf.
3. In C3 plant the efficiency of RuBP carboxylase enzyme is increased
4. Increased CO2 concentration inhibits photorespiration in plants
5. CO2 enrichment increased the yield and yield components.
Other green house gases
1. Oxides of nitrogen (NO, NO2, N2O molecular N2) cause phototoxic, bleaching and
necrosis (drying of tissues) in plants.
2. Ozone (O3) causes ozone injury to the plants.
Remedial measures for green house effect

Reduction in the use of fossil fuel

Use of alternative sources of energy (renewable energy)

Afforestation and community forestry

Avoiding the use of CFCs and nuclear explosions

Environmental awareness
Global Agriculture to increasing CO2
In 1984, the global average of CO2 is 343 ppm; conc. varies with season &

latitude
Direct effect of CO2 increase in the absence of climatic change

179

Doubling of CO2 from 340 to 680 ppm increases 0 10 % increase in yield of C4

plants(maize, sorghum, sugarcane) and 10 50 % increase in yield in C3 plants
(wheat, soybean, rice) depending upon specific crop and growing conditions

Greater yield benefit accrue to the regions where the C3 rather than C4 crops
dominate

Higher CO2 conc. reduces stomatal aperture, thereby reducing transpiration and
WUE

Doubling of CO2 will cause about 40 % decrease in stomatal conductance in short

term

Law of limiting factor:

When other environmental factors such as water, light, minerals & temperature
limit yield, then higher conc. of CO2 will have little or no effect.

This generalizing concept has been challenged

In certain stressful environments, the relative photosynthesis increased with

increased in CO2 conc.

C3 95 % of worlds biomass is of the C3 category

In C3, O2 compete with CO2 for the site of Rubisco In C4, O2 is not compete with
CO2 for the site of PEPCase

At 340 ppm CO2, in the absence of O2 Rubisco operating only at to of its

substrate saturated capacity

PEPCase has high affinity for CO2 than Rubisco PEPCase is close to CO2
saturation at the present atmospheric CO2 conc., no significant enhance of C4
crop growth from increased CO2 so far as PEP Case is concerned

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Lecture No: 33
SEED GERMINATION - PHYSIOLOGICAL CHANGES DURING SEED
GERMINATION
The process of seed germination starts with the imbibition of water by seed coat
and emergence of growing root tip of embryo. The process ends with the development of
embryo into a seedling.
Physiological and biochemical changes during seed germination
1. Water uptake
Seed germination starts with the imbibition of water by dry seed coat. Due to
imbibition of water, the seed coats become 1) More permeable to O2 and water and 2) less
resistant to outward growth of embryo.
2. Respiration
Rapid increase in respiration rate of embryo occurs. Sucrose is probably the
respiratory substrate at this stage which is provided by endosperm.
3. Mobilization of reserve materials
As germination progresses, there is mobilization of reserve materials to provide.
1. building blocks for the development of embryo
2. energy for the biosynthetic process and
3. nucleic acids for control of protein synthesis and embryonic development
Changes in these components are as follows
i) Nucleic acids
In monocots, during imbibition, there is a rapid decrease of DNA and RNA
content in the endosperm with a simultaneous increase in the embryonic axis probably
due to their transportation as such. High concentration of RNA in the embryonic axis
precedes cell division. Due to more cell division, DNA content is increased.

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ii) Carbohydrates
Insoluble carbohydrates like starch are the important reserve food of cereals in the
endosperm. During germination, starch is hydrolyzed first into maltose in the presence of
-amylase and - amylase and then maltose is converted into glucose by maltase. The
glucose is further converted into soluble sucrose and transported to growing embryonic
axis. During germination, the embryonic axis secretes gibberellic acid into the aleurone
layer which causes synthesis of -amylase.

3. Lipids
Plants like castor bean, peanut etc., store large amount of neutral lipids or fats as
reserve food in their seeds. During germination, the fats are hydrolyzed into fatty acids
and glycerol by lipase enzyme. Fatty acids are further converted into acetyl CoA by the
process, - oxidation. The acetyl CoA is further converted into sucrose via glyoxylate
cycle and is transported to the growing embryonic axis.

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4. Proteins
Some plants store proteins as reserve food in their seeds in the form of aleurone
grains. Proteins are hydrolyzed into amino acids by peptidase enzyme. The amino acids
may either provide energy by oxidation after deamination (removal of amino group) or
may be utilized in the synthesis of new proteins.
5. Inorganic materials
A number of inorganic materials such as phosphate, calcium, magnesium and
potassium are also stored in seeds in the form of phytin. These stored materials are
liberated during germination due to the activity of various phosphatases including
phytase.
Emergence of seedling out of the seed coat
All the changes described above gradually result in splitting of seed coat and
emergence of the growing seedling. The radical comes out first and grows downward,
and then plumule comes out and grows upward. Due to the continued growth of this
seedling, the plumule comes out of the soil, exposed to light and develops its own
photosynthetic apparatus.
Splitting of seed coat may take place either by imbibition pressure or by internal
pressure created by the growing primary root or by hydrolytic enzymes which act on cell
wall contents of seed coat and digest it (e.g. cellulose and pectinase). Sometimes the seed
coat may be extensively rotted by the activity of micro-organisms in the soil.
DORMANCY OF SEEDS
All the viable seeds have capacity to germinate if placed under suitable conditions
necessary for germination. But, some seeds fail to germinate sometimes even if placed
under the condition favourable for germination. This may be due to some internal factors
or due to specific requirement for some environmental factors. During this period, the
growth of the seed remains suspended and they are said to be in rest stage or dormant
stage and this phenomenon is called as dormancy of seeds.

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Factors causing dormancy of seeds

1. Seed coats impermeable to water
The seeds of certain plants especially those belonging to the familys
leguminaceae, solanaceae, malvaceae, etc. have very hard seed coats which are
impermeable to water. The seeds remain dormant until the impermeable layer decay by
the action of soil micro-organisms.
2. Seeds coats impermeable to oxygen
In many plants such as cocklebur and many grasses, the seed dormancy is due to
the impermeability of the seed coat to oxygen. However, during the period of dormancy,
the seed coat gradually becomes more permeable to oxygen so that they may germinate.
3. Immaturity of the Embryo
In certain orchids, the seed dormancy is due to the immaturity of the embryos
which fail to develop fully by the time the seeds are shed. In such cases, the seeds
germinate only after a period or rest during which the development of embryo inside the
seed is completed.
4. Germination Inhibitors
In certain seeds, the dormancy of the seeds is due to the presence of certain
germination inhibitors like coumarin, ferulic acid, abscissic acid, etc. These may be
present in endosperm, embryo, testa or juice or pulp of fruit.
5. Chilling or low temperature requirement
In certain plants such as apple, rose, peach etc, the seeds remain dormant after
harvest in the autumn as they have a low temperature or chilling requirement for
germination. In nature, this requirement is fulfilled by the winter temperatures. In such
case the seeds remain dormant throughout the winter season and germinate only in the
following spring.

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6. Light sensitive seeds

In many species, the germination of the seeds is affected by light resulting in seed
dormancy. Such light sensitive seeds are called photo blastic. Seeds of lettuce, tomato
and tobacco are positively photo blastic and germinate only after they have been exposed
to light. On the other hand, the seeds of certain plants are negatively photo blastic and
their germination is inhibited by light.
Advantages of dormancy
1. In temperature zones, the dormancy of seeds helps the plants to tide over the
severe colds which may be injurious for their vegetative and reproductive growth.
2. In tropical regions, the dormancy of seeds resulting from their impermeable seed
coats ensures good chances of survival.
3. Dormancy of seeds in many cereals is of utmost importance to mankind. If these
seeds germinate immediately after harvest in the field, they will become useless to
man for consumption as food.

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Lecture No: 34
ABSCISSION - SENESCENCE - TYPES, CAUSES, PHYSIOLOGICAL
AND BIOCHEMICAL CHANGES AND REGULATION

SENESCENCE AND ABSCISSION

Like human beings, plants a1so grow old and undergo aging and then they die.
Aging is the sum total of changes in the total plant or its organs. During aging, the plants
undergo chemical and structural changes. Aging leads to senescence and later phase of
development that u1timately terminates to death.
Senescence
The deteriorative process which naturally terminates the functional life of an
organ, organism or other life unit is collectively called senescence. Senescence is a phase
of the aging process. The major characteristic of senescence is that the metabolic
processes are catabolic and eventually become irreversible and terminate to death.
Senescence is not confined only to whole plant. It may be limited to a particular
plant organ such as leaf and flowers or cells or cell, organelles. Senescence is closely
associated with the phenomenon of aging. Aging leads to senescence. Wheat plant dies
after the development of fruit. This is the senescence of an entire plant. Leaf fall in a
coconut tree is an example of senescence.
Types of senescence
Leopold (1961) has proposed types of senescence patterns in plants which are as follows.
(a) Overall Senescence
This type of senescence occurs in annuals where whole plant is affected. It is also
called whole plant senescence. The entire plant dies after the development of fruit and
seeds. E.g. Paddy, wheat, soybean etc.
(b) Top Senescence

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In top senescence, the parts remaining above the ground or (shoot system) may die,
but the root system and underground system remain viable. It is also called shoot
senescence. E.g. Dock, perennial herbs.
(c) Deciduous Senescence
In deciduous woody plants, all the leaves die but the bulk of the stem and root
system remains viable. It is called deciduous senescence or simultaneous or synchronous
senescence. E.g. Leaf fall in deciduous trees.
(d) Progressive Senescence
It is a gradual death of old leaves from the base to the top of the plants. It may
occur at any time. It is also called sequential senescence. E.g. Leaf fall in a coconut tree.
Causes of Senescence
1. Leaf senescence is accompanied by early loss in chlorophyll, RNA and enzymes.
2. Cellular constituents are decreased due to slower synthesis or faster break down.
3. Competition between vegetative and reproductive organs for nutrients.
4. A senescence factor (a hormone) is produced in soybean fruits that move to leaves
where it causes senescence.
5. Short-day and long-night conditions induce flowering and leaf senescence.
6. Degradation of food reserves and loss of integrity in food storage cells of seeds.
7. Senescence is also hormonally controlled.
Physiology of Senescence
The following physiological changes occur during senescence.
1. Photosynthesis stops.
2. Chlorophyll degradation: The colour of leaf changes from green to yellow.
3. Anthocyanin pigments accumulation in the leaves causing reddening in leaves.
4. The vacuoles function as lysosomes and digest the cellular materials.
5. The starch content decreased.
6. RNA and proteins are decreased.

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7. DNA molecules are degraded by the enzyme DNase.

8. Growth promoting hormones such as cytokinin decrease.
9. The deteriorative hormones such as ethylene and abscisic acid (ABA) content are
increased.
Senescence Promoters
Senescence is promoted by hormones such as abscisic acid and ethylene. The
senescence accelerating ability of abscisic acid is well documented. The function of ABA
as a promoter of flower tissue senescence including initiaton of colour fading or blueing
has been established. The ABA content of aging leaves increases markedly as senescence
is initiated. Ethylene plays a very important role in the senescence of certain plant parts,
particularly fruit and petals and in the abscission process. It is an inducer in the
senescence of flower tissue.
Senescence Retardants: The primary plant hormones involved here are auxin, gibberellin
and cytokinin.
Significance of Senescence
1. The whole plant senescence occurs in monocarpic plants coinciding the seed
setting and seed dispersal.
2. Due to the formation of abscission layer, the older leaves tend to fall down so that
the nutrients will be diverted to the next young leaf.
3. The senescence process helps the mobilization of nutrients and of the vegetative
parts of the plant into the fruits.
4. Plants escape the influence of seasonal adversity by undergoing senescence of its
organs. Leaf fall in deciduous trees reduces the rate of transpiration to survive
under adverse conditions.
Abscission
Shedding of leaves, flowers and fruits is called abscission. Abscission is distinct
in deciduous trees and shrubs. In autumn, all the leaves of deciduous plants fall, at about
the same time giving the plants a naked appearance. In evergreen plants there is gradual
abscission of leaves. The older leaves fall while new leaves are developed continuously
throughout the year. In most of the herbaceous species, however the leaves are not shed

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even after they die. In many cases leaves are retained in withered dry condition even after
the whole shoot is dead.
Abscission is a complex physiological process. During abscission, the colour of the
leaves, flowers and fruits changes due to degradation of chlorophyll and the synthesis of
anthocyanin pigment.
Leaf abscission takes place at the base of the petiole. The site of abscission is
internally marked by a distinct zone called abscission zone. This zone is made up of one
or more layers of cells arranged transversely across the petiole base. This is called
abscission layer. The abscission zone is pale or brown in colour. The cells of the
abscission layer separate from each other due to the dissolution of middle lamellae and
the primary cellulose walls under the influence of the activity of enzymes, pectinase and
cellulase.
At this stage, the petiole remains attached to the stem by vascular elements only. But
due to its own weight and the wind force, the leaf is detached from the stem. The broken
vascular elements are soon plugged with tyloses or gums. Wound healing in cells
proximal to the breaking point involves formation of a corky layer that protects the plant
from pathogen invasion and excess water loss. Suberin and lignin are synthesized during
healing.
Several environmental factors such as drought and N deficiency promote abscission.
Auxin is synthesized in growing leaf blades and it strongly retards senescence and
abscission. Abscission starts when the amount of auxin begins to decrease. Cytokinins
and gibberellins arriving from the roots also delay senescence and abscission. Abscission
is caused by the formation of cell wall degrading enzymes in the abscission zone, due to
ethylene production.
Significance of Abscission
1. It helps in diverting water and nutrients to the young leaves
2. It is a self pruning process through which fruits and injured organs are shed from
the parent plant.
3. It helps in disseminating fruits and vegetative propagates.
4. Abscission serves as function in removing plant parts containing waste materials.

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