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AGY 202 NOTE

1.0 PLANT WATER RELATIONS
1.1 WATER

Water is an indispensable component of life. It is an important part of all living things some of
which may contain more than 90 percent. In green plants, water account for between 80 to 95 %
of the fresh weight, especially, the woody species, most of which are contained in the vacuoles of
the cells. Here, it provides a suitable medium for many biochemical reactions. Apart from this,
water play many other important roles in the physiology of plant because of its uniquely fitted
physical and chemical properties.

1.1.1 Structure of Water

Water is made up of two atoms of hydrogen joined to one of oxygen atom by covalent bonds
forming an angle of 105°. Since the oxygen atom is more electronegative than the hydrogen atom,
the electrons of the covalent bonds tend to be attracted towards oxygen atom resulting in partial
negative charge on oxygen and equal partial positive charges on each of the hydrogen in water
molecule. As a result of the equal partial negative and positive charges, water molecule carries no
net charge and is neutral. The partial negative and positive charges on two sides of water molecule
make it a polar moleculewith the result that positive side of one water molecule is attracted to the
negative side of another water molecule and joined by hydrogen bond, usually represented by
dotted line. These bonds provide water with unique physical properties.
Hydrogen bond is a weak electrostatic chemical bond formed between covalently bonded
hydrogen atom and an electronegative atom with a lone pair of electrons such as nitrogen or oxygen
and is represented by dotted line. The energy of hydrogen bond is lesser than ionic or covalent
bond but higher than Van der Waals forces and it varies from 8 – 42 kJ mol-1 of bonds. In plants,
hydrogen bonds may also be formed between water and other substances especially those which
contain electronegative O and N atom with lone pairs of electrons.

1.1.2. Properties of Water

Pure water is a colourless, odourless liquid with the molecular weight of 18 Dalton, melting point
of 0°C, boiling point of 100°C and maximum density of 1 gcm-3 at 4°C. Water is liquid at room
temperature. The physical properties of water include:

a. Specific Heat: This is the amount of heat energy required to raise the temperature of a unit
mass of a substance by 1°C. For 1 g of water, the value is 4.2 J. Thus, the specific heat of
water is 4.2 J g-1. This value is higher than that of other liquids, except liquid ammonia.
This property is as a result of the hydrogen bonds between water molecules. When the
temperature of water is raised, the molecules vibrate faster and absorb large quantities of
energy to break the hydrogen bonds thus requiring relatively large amount of energy to
raise its temperature compared to other liquids. This property of water is important to plants
in protecting them from potentially harmful temperature fluctuations.
b. Latent Heat of Vaporization: This is the energy required to convert liquid into gas
(vapour) phase at constant temperature. Water has a latent heat of vaporization of 2.4 kJ g-
1
or 44 kJ mol-1 at 25°C which is the highest known value among all liquids. The higher
latent heat of vaporization of water gives cooling effect to plants as they dissipate heat
through foliar respiration.

c. Latent Heat of Fusion/Melting: It is the heat energy needed to convert a unit mass of a
solid to liquid at constant temperature. For instance, to melt 1 g of ice at 0°C, 333.6 J of
energy will be required. Thus water has a latent heat of fusion of 333.6 Jg-1. In ice, each
water molecule is joined to four others by hydrogen bonds forming a tetrahedral structure,
whereas in water, each molecule of water is joined to five or more. The tetrahedrons are
then arranged in such a way that ice crystals are basically hexagonal. When ice melts to
liquid water, the water molecules move farther apart. However, it is noteworthy that the
volume actually decreases during melting.

O
H

O
H
H
H
O
O
O

d. Water Expansion and Density: Water has the tendency to expand as it freezes and its
density is decreased.

e. Cohesive and Adhesive Force: Mutual force of attraction between like molecules such as
in water (due to H-bond) is referred to as cohesion. On the other hand, attraction of water
to a solid surface such as cell wall or glass surface is called adhesion. Cohesive and
adhesive properties of water are of great importance in ascent of sap in plants.
 f. Surface Tension: Water has higher surface tension than most other liquids (except
hydrazine and most metals in liquid state such as mercury). The high surface tension
enables water to move easily by capillarity through micro-spaces between soil particles
and the plant cell walls.

g. Tensile Strength: This is the ability to resist pulling without breaking, usually measured
as force per unit area. The cohesive property of water gives it a high tensile strength which
enables water in the xylem to be pulled to the top of the plant without breaking.

h. Viscosity: This property of water influences how fast it flows in response to pressure
gradient in the plant.

i. Water as a Solvent: The small size of water molecules and its polarity makes it an
excellent solvent for three groups of biologically important solutes, namely;

i. Organic Solutes with which water can form hydrogen bonds such as amino acids, low
molecular weight carbohydrates and proteins.

ii. Charged ions such as the major nutrient ions (K+, Ca2+, H2PO4-, NO3-, etc.).

iii. Small molecules, such as atmospheric gases (O2, N2) which presumably can fit into
voids in the rather open structure of liquid water.

This property of water makes it an excellent medium for the transportation of organic
substances (e.g. sucrose in the phloem), inorganic ions (e.g. nutrients from the root to the
leaf in the xylem) and atmospheric gases (e.g. diffusion of oxygen to sites of respiration).

j. Light Absorption by Water: The colourless nature of water enables the penetration of
visible light. This property is beneficial to aquatic plants which enable them to
photosynthesize even when they are buried at considerable depths in the water body. Water
also acts as heat insulator since it absorbs the infra-red light. Long wavelength thermal
radiation is also absorbed by water vapour in the atmosphere.

1.1.3. Importance of Water to Plant

Water constitutes 80 – 95% of the total weight of growing plant tissues. Seeds which are the driest
plant tissues, still contain 5 – 15% of water, and must absorb considerable amount of water before
they can germinate. Thus plant play important role in the life of plants.

a. Water is the best know solvent and thus provides medium for the movement of
molecules within and in between the cells

b. Almost all molecules of protoplasm owe their specific biochemical activities to water
environment in which they exist.

c. The structures of macromolecules such as proteins, nucleic acids, polysaccharides and

other cell constituents are greatly influenced by water

d. Water takes direct part in many biochemical reactions in the cell such as hydrolysis,
hydration, dehydration, etc. Water is also one of the raw materials in photosynthesis.

e. Through transpiration, water plays an important role in controlling the temperature of

plants.

f. Plant cells have large central vacuole filled with cell sap and develop large intracellular
pressure called tugor pressure which is essential for many physiological processes in
plants such as (i) cell enlargement, (ii) stomatal movements, (iii) transport of solutes in
phloem, (iv) transport processes in cell membranes, (v) maintaining shape or form of
plant tissues, (vi) emergence of young seedling from soil, etc.

g. Water is the most important factor for agricultural productivity.

h. Water is an essential factor in completing the lifecycles of lower forms of plant life and
aquatic higher plants.
1.2 PERMEABILITY, DIFFUSION, OSMOSIS AND IMBIBITION

1.2.1 PERMEABILITY

Permeability is the ability of a membrane to allow the passage of gases, liquids and liquid dissolved
substances through it. There are two types of membrane:

i. Permeable Membranes: These are membranes that allow all solutes and solvents to
pass through them. Examples are cellulosic cell walls and lignified cell walls.
Permeable membranes are of three types:
 fully permeable allow the passage of both solutes and solvent (e.g. cellulosic cell
wall);
 semi-permeable allows the passage of only solvent molecules (e.g. colloidal
membrane); and
 selectivelyor differentially permeable allows all solvent molecules and selected
dissolved solutes to pass through them (e.g. plasma membrane).

ii. Impermeable Membranes: these are membranes that do not allow solutes and
solvents to pass through them (e.g. cutinized cell wall).

1.2.1 DIFFUSION

Diffusion, a word coined by R. Brown in 1928, is the movement of particles or molecules from
region of higher concentration to the region of lower concentration. The rate of diffusion of gas is
usually faster than that of liquid. Diffusing particles or molecules have a certain pressure known
as the diffusion pressure which is directly proportional to the number of concentration of the
diffusing particles. This is why diffusion always takes place from a region of higher diffusion
pressure to a region of lower diffusion pressure, i.e., along diffusion gradient.

j. Factors Affecting Diffusion

a. Density of the Diffusing Particles: the rate of diffusion is inversely proportional to the
square root of the density of the diffusing particle. Hence, heavy molecules diffuse slowly.

1
𝑟∝
√𝑑

Where 𝑑 = density of diffusing particles

 𝑟 =rate of diffusion

b. Resistance of the Medium: Diffusion rate is inversely proportional to the frictional

resistance of the medium in which diffusion is occurring. Thus, the higher the frictional
resistance of the medium, which is dependent on the density, the lower the diffusion rate.

1
𝑟∝
𝑓

Where 𝑓 = frictional resistance of the medium

𝑟 =rate of diffusion

c. Temperature of the System: Rate of diffusion is directly proportional to the temperature

of the diffusing particles as temperature increases the kinetic energy of the particles.
d. Diffusion Pressure Gradient: Rate of diffusion is directly proportional to the diffusion
pressure gradient; the higher the gradient, the higher the diffusion rate.

ii. Significance of Diffusion

a. Diffusion is an effective means of transport of substance over a short distance. It helps in

phloem translocation, absorption of water and minerals.
b. The process is significant in transpiration process
c. Transfusion of gases such as CO2 and O2 occurs through stomata by this process
d. Ions and hormones move from one cell to another by this process

1.2.2 OSMOSIS

Osmosis is the diffusion of molecules of solvent into a solution through a semipermeable

membrane. Or when a solution is separated from its weaker solution by a semipermeable
membrane, the weaker solution passes into the stronger solution so as to equalize concentration on
both sides. It is the movement of solvent molecules, from the region of higher concentration to the
region of lower concentration, through a semipermeable membrane.

i. Osmotic Potential
When a solution of two different concentrations is separated by a semi-permeable membrane, a
pressure is developed in the solution with higher concentration. It is the pressure required to stop
the entry of the solvent molecules, and is called Osmotic Pressure(OP). It is measured in
atmospheres. More concentrated solution has higher osmotic potential. Thus in osmosis,
movement of solvent molecules takes place from the solution with lower OP (i.e. less concentrated
or hypotonic) to the solution with higher OP (i.e. more concentrated or hypertonic). Osmosis does
not occur in isotonic solutions (equal concentrations).

ii. Plant Cells as Osmotic Systems

Living cells in plants form osmotic system due to the presence of plasma membrane, a
semipermeable membrane, with the cell sap having a certain osmotic pressure. The solvent in
plants is water. The cell wall is permeable.

iii. Endosmosis

If a living plant cell or tissue is placed in water or hypotonic solution, water enters into the cell sap
by osmosis. The process is called Endosmosis. The entry of water into the cell sap causes a
pressure to develop pressing the protoplasm against the cell wall and the cell becomes turgid. The
pressure that develops as a result of endosmosis is called turgor pressure (TP). The turgor pressure
also leads to equal and opposite pressure exerted by the elastic cell wall called the wall pressure
(WP).

𝑇𝑃 = 𝑊𝑃

iv. Exosmosis

In exosmosis, the direction of water movement is to the outside. When a cell is placed in hypertonic
solution, water comes out of the cell sap into the outer solution and the cell becomes flaccid. If the
situation continues, the protoplasm becomes separated from the cell wall and assumes a spherical
shape. This phenomenon is called plasmolysis and the cell is said to be plasmolysed. Because of
the permeable nature of the cell wall, the space between the plasma membrane and the cell wall
becomes filled with the hypertonic solution.

If a plasmolysed cell or tissue is placed in hypotonic solution, endosmosis occurs and the
protoplasm assumes its normal shape and position. The phenomenon is called deplasmolysis.
v. Significance of Osmosis

a. It plays a vital role in water absorption from soil

b. Cell to cell transport of water occurs in plant bodies by this process
c. Shape and form of organisms, turgidity and expansion of a leaf is maintained due to
osmosis
d. Osmosis causes turgidity of young root cells which helps in penetration of soil particles.
e. Stomatal movement is controlled by this process
f. Osmosis plays a vital role in bursting of many fruits, folding and unfolding of leaves of
Mimosa plants.

vi. Diffusion Pressure Deficit (DPD) or Suction Pressure (SP)

The difference between the diffusion pressure of the solution and its solvent at a particular
temperature is termed diffusion pressure deficit (DPD). It was a term coined by B. S. Meyer in
1938, while Renner (1915) coined the term suction pressure. Now the term water potential is used
in place of DPD but with opposite sign. The DPD of a cell sap is a measure of the ability of the
cells to absorb water. It is related with OP and TP of cell sap and also the wall pressure (WP) as
follows:

𝐷𝑃𝐷 = 𝑂𝑃 − 𝑊𝑃

𝑊𝑃 = 𝑇𝑃

𝐷𝑃𝐷 = 𝑂𝑃 − 𝑇𝑃

vii. DPD at Different Cell Conditions

a. At Turgid State:
𝐷𝑃𝐷 = 𝑂𝑃 − 𝑇𝑃 (𝑊𝑃 = 𝑇𝑃)
𝑂𝑃 = 𝑇𝑃 (𝑖𝑛 𝑓𝑢𝑙𝑙𝑦 𝑡𝑢𝑟𝑔𝑖𝑑 𝑐𝑒𝑙𝑙)
𝐷𝑃𝐷 = 𝑂𝑃 − 𝑇𝑃 = 0
Cell has no capacity to absorb water.
b. At flaccid State:
𝐷𝑃𝐷 = 𝑂𝑃 − 𝑇𝑃
𝑇𝑃 = 0 (𝑖𝑛 𝑓𝑢𝑙𝑙𝑦 𝑝𝑙𝑎𝑠𝑚𝑜𝑙𝑦𝑠𝑒𝑑 𝑐𝑒𝑙𝑙)
 𝐷𝑃𝐷 = 𝑂𝑃
Cell absorb water at a force equal to that of the OP

viii. Concept of Water Potential, Solute Potential and Pressure Potential

The term Water Potential was coined by Slatyer and Taylor (1960). It is the term used in place of
DPD by recent physiologists. It is equal in magnitude to DPD but opposite in sign. The water
potential of pure water is zero at one atmosphere and a particular pressure, which is the maximum.
The water potential of a solution is less than zero (i.e. negative value). It is represented by Psi
(𝜳𝒘 ). It is expressed in pressure unit. The SI unit for pressure is pascals (Pa).

1 MPa (106 Pa) = 10 bars or 9.87 atmosphere

Solute Potentialis the amount by which water potential is reduced due to the presence of solute.
Osmotic pressure and osmotic potential are numerically equal but the latter has a negative value.
It is represented as 𝜳𝒔 . If 𝑂𝑃 = 20 atms, 𝜳𝒔 = −20 atms.

Pressure Potential is the potential created by the wall and turgor pressure of plant cells. Unlike
the first two mentioned earlier, pressure potential has a positive value and is represented as 𝜳𝒑 .

𝜳𝒘 = 𝜳𝒔 + 𝜳 𝒑 + 𝜳𝒈

Where 𝜳𝒈 is the gravitational potential which denotes the effect of gravity on the water potential
of a water column in a vertically growing plant. It is usually negligible in plants shorter than 5 m
and thus not taken into account. Therefore the prevalent equation is:

𝜳𝒘 = 𝜳𝒔 + 𝜳𝒑

If for example, there are 2 cells A and B in contact with each other. The cell A has a pressure
potential of 4 bars and contains sap with osmotic potential of – 12 bars. Cell B has pressure
potential of 2 bars and contains sap with osmotic potential of -5 bars. Then,

𝜳𝒘 𝑜𝑓 𝑐𝑒𝑙𝑙 𝐴 = 𝜳𝒔 + 𝜳𝒑

= −12 + (+4) = −8 𝑏𝑎𝑟𝑠

𝜳𝒘 𝑜𝑓 𝑐𝑒𝑙𝑙 𝐵 = 𝜳𝒔 + 𝜳𝒑
 = −5 + (+2) = −3 𝑏𝑎𝑟𝑠

1.2.3 IMBIBITION

It is a physical process that involves the adsorption of water molecules or any other liquid
molecules by hydrophilic substances in the cell, making them swell. Such substances include
cellulose, protoplasm, starch, proteins, etc. Molecules which imbibe water are termed imbibants.
Oily seeds contain more hydrophilic substances than proteinaceous and starchy seeds. In
imbibition, volume is increased, heat is released, known as heat of wetting, and pressure is
developed termed imbibitional pressure or matrix potential, represented as 𝜳𝒎 . It is related to
water potential by:

𝜳𝒎 = 𝜳𝒘 + 𝜳𝒑

If the imbibant is no confined, no turgor pressure is involved and hence the above equation is
simplified to:

𝜳𝒘 = 𝜳 𝒎

Imbibition plays a very important role in the life of the plants. Imbibition initiates seed
germination. 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.
1.3 ABSORPTION AND TRANSLOCATION OFWATER

1.3.1 Soil Water

Soil water is the water present in the soil and it varies in quantity, ranging from traces to abundance.
The soil receives its water from diverse sources such as rains, irrigation of fields, melting of snow,
etc. However, not all the water received by the soil at a particular time are held in the soil, and the
water held in the soil is of different types. They include gravitational water, runaway water,
hygroscopic water, capillary water and chemically combined water.

i. Gravitational Water: This is the water that moves under the influence of gravitational
force. After rains or irrigation of the field, water remains temporarily on the surface of the
soil and gradually percolates deep and deep into the layers of the soil. The air in between
the soil particles are replaced by water molecules. Ultimately, the water goes down below
the reach of the roots of the plants and thus not available to the plants for absorption. At a
point, the soil layers become saturated, forming the water table. Immediately after rains or
irrigation, the water is temporarily available around the root zone for absorption by the
plants. Logging occurs when rain water remains around the root zone for a long time, and
it causes injury to the plant root by preventing the penetration of oxygen and causing an
accumulation of excess carbon dioxide around the root zone.

ii. Run-away Water: After the surface of the soil is saturated, the majority of rain water gets
drained off horizontally. The flow of water on the surface is termed runaway water or run
off water. Runaway water differs from gravitational water in that, while the former flows
horizontally, the latter flows vertically under the influence of gravitational force.

iii. Hygroscopic or Imbibed Water: This is the water held by colloidal soil particles by
adsorptive forces between the soil particles and water molecules. Due to these forces, the
osmotic potential of hygroscopic water is greatly reduced; hence it is not available to plants.

iv. Capillary Water: This is the water found between the spaces of non-colloidal soil particles
in the form of a thin film. This water is held under the influence of surface tension or
capillary force and is most suited for absorption by roots of plants. The finer the texture of
the soil, the greater the capillary water held.
 v. Chemically Combined Water: Some of the minerals like silicon, iron, and aluminum have
water in the combined form. This water is not available to plants for absorption. Chemically
combined water gets evaporated on heating at 105°C. After hygroscopic water is gone from
the soil, the only water that remains in the soil is in hydrates of aluminum, iron, silicon,
etc.

 The total amount of water present in the soil is called holard

 Water that is available to plants is called chesard
 Unavailable water is called echard. Thus, holard is the sum of chesard and echard.
 Field Capacity: this is the amount of water retained by the soil after rains or irrigation,
against the force of gravity. It is also called water holding capacity of the soil.
 Permanent Wilting Percentage/Wilting Coefficient: this is the percentage of soil water
that remains in the soil when the plants have shown permanent wilting. It usually ranges
from 1% to 15%of water. Although it is dependent soil type, it has been shown that for a
particular type of soil, the value differ from one plant to another.

1.3.2 Water Absorption

In higher plants, water is absorbed through root hairs which are in contact with soil water and form
a root hair zone a little above the root tip. Root hairs are tubular hair like prolongation of the cells
of the epidermal layer of the roots. The walls of the root hairs are permeable, and consist of pectic
substances which are strongly hydrophilic. They contain vacuoles filled with cell sap with osmotic
pressure of between 3 – 5 atms (that of the outer soil water is usually less than 1). When roots
elongate, the older hairs die and new ones are formed.

i. Mechanism of Water Absorption

There are two independent mechanisms for water absorption in plants. They are:

a. Active Absorption of Water: this is when the root hairs play active role in the
absorption of water and the metabolic energy released through respiration is
consumed. It may be of two types:
 Osmotic Absorption: the first step here is imbibition of water by the hydrophilic
cell walls of the root hairs. The OP of the cell sap of the root hair is usually
higher than that of the soil water thus creating a higher DPD, and water from the
 cell walls enters into them through the plasma membrane by endosmosis. As a
result, the OP, DPD and the SP of the root hair become lower while their turgor
pressure is increased. Water from the root hairs now moves to the inner cells of
the root along concentration gradient and finally reaches the xylem where it goes
up to the leaves for consumption.
 Non-Osmotic Absorption: It has been observed that absorption sometimes takes
place even when the OP of the soil is greater than that of the cell sap. This type
of absorption, which is non-osmotic and against the osmotic gradient, involves
the expense of metabolic energy, probably supplied through respiration of the
cells. It has been observed that the factors which inhibit respiration also decrease
water absorption.
b. Passive Absorption of Water: This occur when the root hairs play passive role in the
absorption of water as the forces responsible for absorption are not generated in the
root but in the upper parts of the plant, i.e. transpiration pull.
ii. Factors Affecting Water Absorption
a. Availability of water: rate of water absorption increases with the increase in soil water
content up to its field capacity.
b. Concentration of the soil water: absorption generally takes place when the
concentration of the cell sap is more than that of the soil water.
c. Soil temperature: absorption of water is decreased at a temperature below 20°C and
at temperature higher than 30°C.
d. Aeration: absorption of water is retarded in poorly aerated soils because such
condition results in deficiency of oxygen and accumulation carbon dioxide, thereby
retarding the metabolic activities of the roots like respiration.

1.3.3 Translocation of Water (Ascent of Sap)

Water after being absorbed by the roots is distributed to all parts of the plant and the excess is lost
through transpiration. In order to reach the topmost parts of the plant, water has to move upward
through the stem. This upward movement of water in plant against gravitational force is known as
ascent of sap.
i. The Path of Ascent of Sap: the path of ascent of sap is the xylem which can be shown
by the following experiments:
a. When a cut shoot of balsam (it has a semi-transparent stem) is immersed is water
containing eosine (a dye), after some time, coloured lines will be seen moving
upward in the stem. A transverse section of the stem will show that only xylem is
stained.
b. Ringing experiment: a leaf 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 ringed part remained fresh and green.
ii. Mechanism of Ascent of Sap: although the mechanism of ascent of sap is not well
understood, a number of theories have been put forward to explain it.
a. Vital Forces Theories: Supporters of these theories think that ascent of sap is under
the control of vital activities in the step.
 Godlewski (1884) proposed that ascent of sap takes place due to the pumping
activity of the cells of xylem parenchyma and medullary ray. It was later
disproved by the experiments of Strasburger (1891, 1893) who demonstrated
that ascent of sap continues even in the stems in which living cells have been
killed by the uptake of poison.
 Bose (1923) gave the pulsatory force theory in which he proposed that the
pulsatory activity of the innermost layer of the cortex (just outside the
endodermis) is responsible for ascent of sap.
 Root Pressure Theory: the pressure developed in the xylem vessel as a result
of the metabolic activities of the roots. Root pressure is developed when rate
of water absorption is more than the rate of transpiration, and as a result, water
is pushed up in the trachery elements of roots.Strasburger has demonstrated
ascent of sap in plants in which the roots are removed.

b. Physical Forces Theories: A number of physical theories have been proposed

which state that ascent of sap occur due to some physical forces like capillary
imbibition, atmospheric pressure, transpiration pull, etc.

 Imbibition theory: It was proposed by Unger (1868) and supported by Sachs

(1878). According to this theory, ascent of sap takes due to imbibitional force
 generated by the thick walls of xylem cells due to the possession of hydrophilic
colloids. Objection: it has been found that water moves through lumen of cell
and not cell walls.
 Capillary theory: Proposed by Boehm (1809). He proposed that tracheids and
vessels resemble capillaries, and so water rises in the narrow tubes of xylem
(tracheid + vessels) due to capillary force. The factors governing the rate of
water movement through capillaries are expressed in Poiseuille’s law
(Poiseuille, 1840).

𝜋𝑟 4 ∆𝑃
𝐽𝑣 =
8 𝑛𝑙

where𝑛 = viscosity,

∆𝑃
−mean pressure gradient along the capillary
1

𝑟 =radius of capillary, 𝑙 = length

The volume of fluid moving in unit time along a cylinder is proportional to the
fourth power of its radius, and the movement depends linearly on the drop in
hydrostatic pressure.

Objection: tracheids and vessels, unlike cylinder, have end walls; vascular
system is not open to the air at the upper end and thus vessels are not analogous
to a capillary.

 Atmospheric pressure: the water transpired from the surface of the leaves
reduces the pressure in the xylem cells and this gap is filled by the water just
below it due to atmospheric pressure; this leads to the uplifting of water in the
stem. Objection: this pressure can only raise water up to 34 feet.
 Transpiration pull and cohesion of water: Proposed by Dixon and Joly (1894)
and supported by Dixon (1914, 1924). It is very convincing and the most
widely accepted. It is based on the following features:
 Cohesive and adhesive properties of water molecules makes it
form a continuous water column in the xylem;
  Transpiration pull exerted on the water column in the xylem.

Water forms a continuous column from root to leaf through xylem ducts. When
water leaves the mesophyll cells of the leaves due to transpiration, the TP
decreases leading to an increase in their DPD (remember, DPD = OP – TP). As
a result, they suck water from adjacent xylem vessels of the leaves and thus
xylem sap is under pull or tension. This pull is transmitted downward and
relieved when water is absorbed through the roots. The column of water resists
breaking due to forces of cohesion and adhesion.
 2.0 TRANSPIRATION, GUTTATION AND EXUDATION

2.1 TRANSPIRATION

Transpiration is the loss of excess water in the form of vapour from the aerial green parts of plants.
These include green leaves, stem and green fruits. Transpiration occur by day as well as by night.
Ninety five percent of transpiration occurs during the day while the remaining five percent occur
by night. During the day, transpiration is maximum at noon between 11 am and 3 pm. The amount
of water lost in transpiration varies from plant to plant.

2.1.1 Types of Transpiration

There are three types of transpiration, depending on the organ of plant involved:

a. Stomata Transpiration: Most of the transpiration takes place through stomata. Stomata are
usually are usually more on the lower side of the leaves. In monocots however, they are
equally distributed on both sides, while in aquatic plants with floating leaves, they are
present on the upper surface. This form of transpiration is more significant and
predominates. It accounts for between 90 – 97% of the entire transpiration in plant.
b. Cuticular Transpiration: Although cuticle is impervious to water, some water may still be
lost through it. It may contribute between 3 – 9% of total transpiration.
c. Lenticular Transpiration: Some water may be lost by woody stems and fruits through
lenticels. This account for about 1% of the total transpiration.

2.1.2 Stomata

The stoma (plural – stomata) is the main avenue for the escape of water vapour from the leaves of
terrestrial plants. The stoma has two small-sized, differentially thickened green epidermal cells
called guard cells. The common walls are not fused but are free from each other for most of the
length. The guard cells may be further surrounded by two or more specialized cells called
subsidiary cells. Guard cells are connected with nearby epidermal cells by plasmodesmata. They
contain a few small chloroplast and small vacuoles. They are of two types in shape; reniform
(kidney-shaped) and dumbbell-shaped. The latter type occur in grasses while the former occur in
all other plants. The outer walls of guard cells are thick while the inner walls are thin.
The different types of stomata based on distribution and position of stomata on the two surfaces
of the leaf are:

 Apple/Mulberry Type: the stomata are located on the under surface only.
 Potato Type: the stomata are more on the lower side of the leaf than on the upper surface.
 Oat Type: stomata occur equally on both surfaces
 Water lily type: stomata are found only on the upper surface
 Potamogeton type: stomata are either absent or functionless. Most submerged aquatic
plants belong to this category.

On the basis of their opening and closing, stomata are of the following types:

 Cereal (Barley) Type:the stomata are opened for a few hours during the day. Common to
cereals.
 Alfalfa Type: the stomata remain open throughout the day and closes at night. Usually
found in thin leaved mesophytes, e.g. pea, bean, etc.
 Potato Type: the stomata remain open both during the day and night. In case of water
shortage, the stomata close for a few hours at specific time of the day, e.g. potato, onion,
plantain, etc.
 Equisetum Type: here, the stomata seldom close, e.g. Sagittaria.

2.1.2.1 Mechanism of Stomata Transpiration

Mechanism of transpiration takes place in three steps:

i. Osmotic diffusion of water in the leaf from xylem to intercellular spaces above the
stomata through the mesophyll cells. When the mesophyll cells draw water from the
xylem, they become turgid and their DPD and OP drops which makes them release
water in the form of vapour into the intercellular spaces close to the stomata by osmotic
diffusion. This increases their OP and DPD and they draw water from the xylem.
ii. Opening and Closing of Stomata (Stomata Movement): Consequent to an increase in
the OP and DPD of the guard cells, osmotic diffusion of water from the surrounding
epidermal cells and mesophyll cells into the guard cells follows. This increases the
turgor pressure (TP) of the guard cells and they become turgid. The turgidity causes
both walls of the guard cells to stretch, but because the outer wall is thin, it is more
stretched than the inner wall, giving a concave shape to the guard cells and the stomata
open. When the guard cells become flaccid, the walls revert back to their shape leading
to a closure of the opening or stomatal pore.
Mechanism of Stomatal Movement
Based on the time of opening, stomatal movement is of two types; photo-active
movement and scoto-active movement. The former is controlled either directly or
indirectly by light while the latter is not controlled by light. Majority of plants show
photo-active movement while only a few show scoto-active movement (e.g. CAM
plants).
Various theories have been proposed to explain the opening of the stomata:
 Glycollate Hypothesis: It was proposed in 1963 by Zelitch. Glycollate is
synthesized in low CO2 and high O2 concentration of the atmosphere. Glycollate is
converted to carbohydrate via the glycine-serine pathway. This raises the OP of the
guard cells, increasing the DPD which leads to the absorption of water from
adjacent mesophyll and subsidiary cells through endosmosis. The absorption results
in turgidity, causing the stomata to open.
 Starch-Sugar Hypothesis: It was first proposed by Lloyd in 1908 and later
elaborated by many workers such as Sayre and Scarth (1923). Guard cells possess
starch. The starch disappears during opening of stomata, especially during the day,
due to conversion to simple sugar. At the time of opening, the pH of the guard cells
increases which stimulates an enzyme that hydrolyses starch to form glucose-1-
phosphate. Glucose-1-phosphate is then changed to glucose-6-phosphate from
which glucose is formed. Glucose in the guard cells raises the OP of the guard cells,
and as a result, they absorb water from adjoining cells, swell up thereby creating
the opening. In the evening, photosynthesis stops, the reverse of the processes takes
place and the stomata pores are closed.
 K+ Ion Pump Hypothesis: It was proposed by Levitt (1974). At the opening of the
stomata, pH of guard cells rises. The raised pH causes hydrolysis of starch to form
phosphoenol pyruvate. The latter combines with CO2 to form malic acid. Malic acid
dissociates to produce malate and H+ ions. H+ ions are released and the guard cells
absorb K+ ions from adjacent epidermal cells. A lot of Cl- ions are also absorbed to
counter balance the K+ ions. Vacuoles present in the guard cells are useful for
 storing solutes.K+ ions and other ions produce an OP required for absorbing water
from adjacent epidermal cells. On endosmosis, the guard cells swell and produce a
pore between them. During closure, pH of guard cells decrease due to the efflux of
H+ ions, malate is changed to malic acid which causes the leakage of K+ and Cl-
ions out of the guard cells. The loss of ions reduces the OP and the guard cells lose
water to adjacent cells leading to loss of turgidity and the closure of the stomata
pores. Malic acid is slowly changed to form starch.
iii. Simple diffusion of water vapour from the intercellular spaces to outer atmosphere
through the open stomata.

2.1.2.2 Factors Affecting Stomatal Movement

The factors affecting the movement of the stomata include:

a. Light:In photoactive plants, light causes the stomata to open

b. Temperature: At proper temperature, stomata can also open at night. High temperature
enhances the activity of hydrolyzing enzyme, and thus causes the stomata to open. But excess
temperature results in the closure of the stomata, even by day, and the effect varies from
plant to plant.
c. pH: Enzymes which are responsible for the conversion of starch to sugar are pH dependent;
they work in alkaline medium. Thus a rise in pH leads to stomata opening.
d. CO2Concentration:Stomata open at low CO2 concentration and close at high CO2
concentration.
e. Water Content of Leaves: plants under water stress close their stomata to reduce
transpiration. It is triggered by the formation of ABA.
f. Hormones:Cytokinins are essential to stomatal opening. ABA also regulates the stomatal
closure mechanism.
g. Mechanical Shock: Stomata close in high wind due to shock created by the wind.
h. Oxygen: Oxygen is necessary for stomatal opening.
i. Mineral Status: NPK deficiency cause sluggish stomatal movement free phosphate
promotes stomatal closure.
2.1.2.3 Stomatal Index

This is the percentage number of stomata as compared to all the epidermal cells in a unit area of
leaf.

𝑆
𝐼= × 100
𝐸+𝑆

Where 𝐼 = Stomatal index

𝑆 = Number of stomata per unit area

𝐸 = Number of epidermal cells per unit area

The stomatal index is independent of the environmental conditions, age of the plant and position
of the leaf. It is constant for species and has great diagnostic importance.

2.1.3 Factors Affecting Rate of Transpiration

There are two groups of factors affecting the rate of transpiration in plants:

i. External Factors

a. Light: Light increases the temperature of the environment and the transpiring surfaces.
It also increases the permeability of the protoplasmic membrane, leading to increase
in transpiration. High light intensity also causes closure of stomata in some plants
leading to reduction in transpiration rate.
b. Temperature: Increase in temperature results in increase in transpiration rate. Increase
in temperature decreases the relative humidity of the air. Temperature affects vapour
pressure gradient. Rise in leaf temperature causes more transpiration, even in humid
air.
c. Humidity of the Air: Rate of transpiration is increased when the relative humity of the
air drops due to the vapour pressure gradient between the inter and external atmosphere
of the plant.
d. Wind:Wind increases the rate of transpiration by quickly removing the saturated air
around the leaves and replacing it with unsaturated air. High speed wind sometimes
reduces transpiration due to closure of stomata and cooling of the leaf surface.
 e. Atmospheric Pressure: Lower atmospheric pressure reduces rate of transpiration due
to rapid evaporation and development of air currents.

ii. Internal Factors

a. Shape, size, structure and orientation of leaf: These greatly influence the rate of
transpiration. Transpiration is directly proportional to the transpiring surface area; the
larger the transpiring area, the greater the rate of transpiration. The shape of leaf also
influences transpirational rate. And transpiration is more in leaves placed
perpendicular to the incident light than leaves placed parallel to it. Thick cuticle
reduces cuticular transpiration.
b. Stomata: The number of stomata per unit area of leaf (i.e., distribution), the position
of the stomata and the size of the stomata pore influence the rate of transpiration. Rate
of transpiration is minimum in scotoactive plants.
c. Water Content of Leaves: Optimum transpiration occur when the leaves have
sufficient water content; low water content of leaves lower the rate of transpiration by
closing the stomata.
d. Root/Shoot Ratio: High root/shoot ratio increases the rate of transpiration. This is the
case in small plants, hence they transpire more per unit of leaf area compared to large
plants with low root/shoot ratio.
e. Diseases: Rate of transpiration is higher in diseased plants compared to healthy ones.

2.1.4 Transpiration Ratio

This is the ratio of total water transpired to the amount of dry matter accumulated by the same
plant. The ratio is variable in plants, and even plants of the same species growing in different
environmental conditions. The transpiration ratio of Sorghum is 200 – 300 units. The ratio is lower
in desert plants. It is minimum in CAM plants. In pine apple, it is 50 units. In C4 plants, it is 100
– 200 units of water for the manufacture of a single unit of dry matter. For mesophytes, under
normal condition, it is 300 – 500 units of water per unit of dry matter.

2.1.5 Advantages of Transpiration

i. Maintenance of Turgidity: Transpiration helps keep the pressure potential of cells below
their osmotic potential, and thus the cells are not fully turgid.
ii. Removal of Excess Water: Transpiration removes excess water absorbed by the plant, which
are not utilized in metabolic processes and growth of the plant.

iii. Cooling Effect: Transpiration was earlier thought to have a cooling effect on transpiring
surfaces and thus protect them from overheating. It has however recently been found that
plants showing negligible transpiration are not overheated.

iv. Regulation of Water Absorption: Absorption of water by plants depends on transpiration;

higher transpiration results in higher absorption rate.
v. Formation of Mechanical Tissues: It has been observed that increase in the rate of
transpiration favours the development of mechanical tissues which give rigidity and streght
to plants
vi. Regulation of Mineral Salt Absorption: Passive absorption of minerals takes place under
the influence of transpiration, but this type of absorption is not common. Salt is absorbed by
active process, and can occur even in the absence of transpiration.
vii. Controls Ascent of Sap: The theory proposed by Dixon and Jolly (1964) assumes that sap
is uplifted under the influence of transpirational pull.

2.1.6 Disadvantages of Transpiration

i. Loss of Water from Plants: 97% of the entire water absorbed by plants is lost to the
atmosphere by the process of transpiration. Only 3% is retained and utilized by the plant in
various metabolic processes

ii. Wilting: Excessive transpiration results in temporary wilting in plants at noon. When
transpiration/absorption ratio is high, plants face internal water stress. During the day,
transpiration is slightly higher than absorption, while at night, the reverse is the case. If high
T/A persist for sometime, the plant undergoes permanent wilting.

iii. Shedding of Leaves: Deecidious trees shed their leaves in the dry season to minimize
transpiration; minimization of transpiration occur at the expense of photosunthesis.

Considering bothe the advantages and disadvantages of transpiration, it can be concluded that it is
essential, though harmful to the plant. Exchage of gases (CO2 and O2) occurs through the stomata.
Transpiration can be minimized by the use of antitranspirant but cannot be avoided. Curtis (1926)
has regarded transpiration as a necessary evil.

2.1.7 Antitranspirants

Antitranspirants are substances which are applied to transpiring surfaces of plants to check
excessive loss of water through transpiration. They reduce water loss without affecting other
metabolic processes of plants such as exchange of gases. Antitranspirants are of two types:

i. Film Forming Compounds: these are relatively inert compounds which can be spread as
film over the surface of the leaves. They reduce cuticular transpiration. Examples are
colourless platics, silicon oil, low viscosity waxes, etc.

ii. Stomatal Inhibitors: these are compounds which directly affect stomatal movement. Several
herbicides, fungicides and growth regulators act as stomatal inhibitors. They can either be
natural or synthetic. Some natural examples include ABA, phenolic acids and aspirin
(acetylsalicilic acid). Phenyl mercuric acetate, a fungicide, when sprayed at 10-4 M
concetration, causes partial closure of stomata for two weeks but have some toxic effects.
ABA and aspririn have less toxic effect but are costly. An ideal antitranspirant is yet to be
discovered.

2.1.8 Measurement of Transpiration

i. Qualitative Method: By cobalt chloride method, it is possible to measure the transpiration

index of a leaf:

𝐿𝑒𝑎𝑓 𝑡𝑒𝑠𝑡 𝑡𝑖𝑚𝑒

𝑇𝐼 =
𝑊𝑎𝑡𝑒𝑟 𝑡𝑒𝑠𝑡 𝑡𝑖𝑚𝑒

ii. Quantitative Method: Rate of transpiration is measured using a potometer.

2.2 GUTTATION
This is a condition where excess water in the leaves escapes through structures called hydathodes
present at the tips of veins of leaves. Water may sometime escape from the stems through the scars
of leaves and lenticels, this type of water loss is called guttation. It normally occurs in the night
but may also occur in the daytime, if the plants are growing in moist soil under humid condition,
especially when absorption is more than transpiration. Guttation liquid is not usually pure water
but contains both dissolved organic and inorganic substances. It is commonly found in herbaceous
like grasses, potato, tomato, etc.

Hydathodes may be active or passive. Active hydathodes are specialized epidermal cells which do
not have a cuticle on the outside and are not connected directly with vascular strands internally.
Passive hydathodes comprise loosely arranged, colourlessparenchymatous cells known as
epitherm. It lies inside the leaf below the vein end.

2.3 EXUDATION OR BLEEDING

This literally applies to gentle release of watery sap from the cut tissues of plants. Noggle and Fritz
(1976) categorized it into four:

i. Exudation due to development of positive root pressure in xylem vessels. Here the exudate
is called xylem sap.

ii. Bleeding due to pressure developed in the surroundings of the injury. Here the exudate is
xylem sap containing 2 – 3% sugar.

iii. The third type of exudation occur as a result of local pressure developed in the stem when
incision is made in the phloem cells. Here the exudate is phloem sap, as in palm wine from
palm tree. The sap contains up to 20% sugar.

iv. The last type of exudation occurs when incision is made in lactiferous ducts, which are
distributed in the cortex, pith and secondary tissues of the stem of some plants. Here, the
exudate is latex in angiosperm and resin in gymnosperm.

ADAPTATIONS TO REDUCE EXCESSIVE WATER LOSS

Anatomical: a thick cuticle and sometimes a multiple epidermis develop to check excessive
evaporation of water. The presence of cutinized hairs, scales, rods, etc., minimizes transpiration to
a greater extent. A dense coating of hairs, or of wax or bloom, on the surface is very efficient in
this respect. Latex also checks transpiration. The stomata may be closed temporarily even in the
day time. In desert plants, the stomata are fewer in number and they remain sunken in pits. Cork
formed in shrubs and trees also act as waterproof covering.

Morphological:leaf area is often very much reduced. In extreme cases, the leaves are modified
into spines. The size of the plant is also often reduced. The leaves may be rolled up or variously
folded exposing minimum surface area for transpiration. They may also assume a drooping or
vertical position to avoid strong sunlight. Deciduous trees shed their leaves in dry season as a
protection against excessive transpiration while leaves of evergreen trees have their leaves well
covered with cuticle.

HEAT TRANSFER

The energy utilized by plants comes from direct solar radiation. When solar radiation strikes a leaf,
some is reflected, some is absorbed and some is transmitted. Absorbed energy is subsequently
dissipated via three main avenues: re-radiation of long wavelengths to nearby surfaces and sky;
conduction and convection of sensible heat (directly measurable or able to be ‘sensed’); and
latent heat exchange (energy consumed to evaporate water and commonly referred to as
transpirational cooling).

A small amount of energy is used in photosynthesis and some stored in the leaves, but these
quantities are so small (usually 2 – 3% of the total). Thus the larger portion of the energy that hits
the plant surface is dissipated, enabling the plant to maintain an appropriate energy balance.

𝑄𝑎 = 𝑄𝑟 + 𝐶 + 𝐿𝐸 + 𝑃

𝑄𝑎 = 𝐴𝑏𝑠𝑜𝑟𝑏𝑒𝑑 𝐸𝑛𝑒𝑟𝑔𝑦 (𝑏𝑜𝑡ℎ 𝑣𝑖𝑠𝑖𝑏𝑙𝑒 𝑎𝑛𝑑 𝑖𝑛𝑓𝑟𝑎 − 𝑟𝑒𝑑)

𝑄𝑟 = 𝑅𝑒𝑟𝑎𝑑𝑖𝑎𝑡𝑒𝑑 𝐸𝑛𝑒𝑟𝑔𝑦

𝐶 = 𝐸𝑛𝑒𝑟𝑔𝑦 𝑙𝑜𝑠𝑡 𝑜𝑟 𝑔𝑎𝑖𝑛𝑒𝑑 𝑏𝑦 𝑐𝑜𝑛𝑣𝑒𝑐𝑡𝑖𝑜𝑛

𝐿𝐸 = 𝐿𝑎𝑡𝑒𝑛𝑡 ℎ𝑒𝑎𝑡 𝑙𝑜𝑠𝑠

 𝑃 = 𝐸𝑛𝑒𝑟𝑔𝑦 𝑢𝑠𝑒𝑑 𝑓𝑜𝑟 𝑝ℎ𝑜𝑡𝑜𝑠𝑦𝑛𝑡ℎ𝑒𝑠𝑖𝑠

Reflection: A considerable fraction of solar radiation incident on leaves is reflected, and that
fraction varies enormously with wavelength. Reflectance shows a modest peak around green
wavelengths, causing plants illuminated with white light to appear green to our eyes. Reflectance
is especially low for blue and red wave-lengths which are absorbed for photosynthesis. Albedo,
which is the measure of the reflective ability of a surface, varies from plant to plant depending on
their habitat and adaptive measures.

Re-radiation: this is the loss of absorbed heat by the leaves to the sky and nearby surfaces. At
night, leaves are often cooled below air temperature by re-radiation. Thus a leaf will have a positive
energy balance in the day time and a negative energy balance in the night due to the loss of heat
to the sky and the surrounding surfaces through re-radiation.

Conduction: This is a mode of sensible heat transfer by means of molecular agitation within a
material without any motion of the material as a whole. It is a flow of heat from a region to the
region of lower heat. In plants, heat can be transferred by conduction from the leaf surface to the
adjacent cooler air by conduction, and from the roots to the aerial parts, and vice versa.

Convection: This refers to the transfer of heat by mass motion of a fluid such as air or water when
the heated fluid is caused to move away from the source of heat, carrying energy with it. It is
usually vertical. If a leaf surface is cooler than the surrounding air, heat will flow by conduction
into the adjacent air, warming it and making it less dense. The air will then rise from the surface
of the leaf and be replaced by cooler denser air, cooling the leaf surface.

Advection is the horizontal movement of fluid such as air or an ocean current. The case of oasis
effect is a typical example of advection. Warm air is transferred from the hot desert to small
isolated vegetation resulting in a rate of transpiration exceeding that caused by the incident
radiation.

Latent Heat Exchange: This is the loss or gain of heat caused by change of state, such as the loss
when water is evaporated or gained when dew condenses. Transpiration constitutes an important
mode of heat loss, and the reverse applies when water vapour condenses onto a leaf as dew or frost.
PHOTOSYNTHESIS

Photosynthesis is an endergonic and anabolic process that involves the building of simple
carbohydrates such as sugars in the green leaf by the chloroplast in the presence of sun light as the
source of energy from carbon dioxide and water absorbed from the atmosphere and the soil
respectively. It is usually accompanied by the release of oxygen. The volume of oxygen evolved
or liberated has been found to be equal to the volume of carbon dioxide absorbed, but it should be
noted that all the oxygen liberated in the process of photosynthesis comes exclusively from water
and not carbon dioxide.

Oxygen produced escapes into the atmosphere through the stomata. The formation of sugars from
carbon dioxide commonly called carbon assimilation is the monopoly of green plants only,
chlorophyll being indispensible in the process. By this process, not only is simple carbohydrate
formed, considerable amount of energy is transformed by the green plants from light energy to
chemical energy and is stored up as such in the organic substances formed.

Raw Materials of Photosynthesis

The basic materials utilized by plants for the process of photo synthesis include:

1. Carbon Dioxide (CO2): Land plants obtain their CO2 from the atmosphere through the
stomata. Hydrophytes get their CO2 supply from aquatic environment as bicarbonates which
are absorbed by these aquatic plants through their general surface.
2. Water: Higher plants make use of water for their carbon fixation, while photosynthetic
bacteria make use of H2S with no evolution of oxygen. This is a proof that the oxygen evolved
during the process of photosynthesis does not come from CO2 but from water.

6CO2 + 12H2S C6H12O6 + 6H2O + 12S

3. Light: Light is the visible part of electromagnetic spectrum. The visible portion of the
spectrum consists of radiations having a wavelength of 390 – 760 nm. And the radiation
utilized by plant known as Photosynthetically Active Radiation (PAR) falls within this range
of radiation wavelength (400 – 700 nm).
Photosynthetic Apparatus - Chloroplasts

Chloroplasts in green plants constitute the photosynthetic apparatus. In higher plants they are
usually discoid or ellipsoidal in shape. Each chloroplast is bounded by two membranes consisting
of lipid bilayer and proteins. The chloroplast is filled with a hydrophilic matrix called stroma, in
which are embedded grana. Each granum is made up of 5 – 25 disc shaped grana lamellae
(collectively called thylakoids) placed on each other like a stack of coins. Some of the grana
lamellae are connected with others by stroma lamellae or fret membranes. Thylakoid membranes
and stroma lamellae are composed of lipid bilayer and proteins.

Chlorophyll and other photosynthetic pigments are found in the form of protein pigment
complexes mainly in the thylakoid membranes of grana.

Photosynthetic Pigments

Photosynthetic pigments are of three types; chlorophyll, carotenoids and phycobilins. Chlorophyll
and carotenoids occur in green plants while phycobilins are found in some algae.

a. Chlorophyll: this is the green pigment in plant with types ranging from chlorophyll a to e
with little structural differences.Of these forms of chlorophyll, chlorophyll a and b’s
structures and functions are well known. Structurally, chlorophylls are generally made up
of a Porphyrin head and a phytol tail
- Chlorophyll a: It has a molecular formula C55H72O5N4Mg and a molecular weight of
893. It absorbs blue, yellow and red wavelengths of light spectrum.
- Chlorophyll b: It has a molecular formula C55H70O6N4Mg and a molecular weight of
907. It is similar to chlorophyll a but with a formyl (CHO) group instead of methyl
 (CH3) group at carbon 3 position of the second pyrol ring. It absorbs blue and red
wavelengths.

b. Carotenoids: These are unsaturated polyhydrocarbons which do not require light for their
biosynthesis. They are of two types; carotene and xanthophylls.

- Carotene: They are orange coloured pigments having the formular C40H56 and a
molecular weight of 536. They are found in all groups of plants and occur as α-carotene
and β-carotene. On hydrolysis, β-carotene produces Vitamin A. It is abundant in
carrot.

- Xanthophylls: They are oxygen containing carotenoids also called carotenols or

xanthols, and are more abundant in nature with the ratio of 2:1 xanthophyll to carotene
in young leaves. The most common xanthophyll in green plant is lutein (C40H56O2).

c. Phycobilins (Biliprotein):Phycobilins consist of a bile pigment attached to a protein.

Three classes are recognized; phycoerythrin (red), phycocyanin (blue) and allophycocyanin
(light blue). They are mainly found in red and blue-green algae.

Photosynthetic Units

These are groups of pigment molecules that take part in the change of light energy into chemical
energy. Each unit has a reaction centre of special chlorophyll a molecule which absorbs light of
long wavelength, and are surrounded by a number of light harvesting pigment molecules. These
light-harvesting pigment molecules are of two types; the antenna and core molecules.

The antenna molecules are pigment molecules that occur on the side of photosynthetic unit. They
absorb photons of different wavelengths, get excited, and they then transfer the energy to core
molecules by electron spin resonance. The core molecules are pigment molecules which lie around
the trap centre. They take part in both direct light harvesting as well as transfer of energy from
antenna molecules. The energy gained by the core molecules is passed on to reaction or trap centre.
With the gain of energy, the reaction centre releases an electron.

Pigment Systems (Photosystems)

Photosynthetic units occur in two distinct groups called the photosystems or pigment systems.
Green plants and cyanobacteria possess two photosystems I and II.

Photosystem I (PS I): This is the photosynthetic system along with some electron carriers that is
located in the non-appressed part of the grana thylakoids as well as the stroma thylakoids. The
photocentre (reaction centre) has a special chlorophyll a molecule called P700. The P700 is
surrounded by other chlorophyll a, followed by chlorophyll b and carotenoids. PS I contains FeS,
ferredoxin, plastoquinone, cytochrome complex and plastocynanin. It takes part in both cyclic and
non-cyclic photophosphorylation, and can carry on cyclic photophosphorylation independently.
PS I drives an electron from PS II to NADP+.

Photosystem II (PS II): It is a photosynthetic pigment system along with some electron carriers
that is located in the appressed part of the grana thylakoids. PS II has chlorophyll a, b and
carotenoids. Its chlorophyll a and b content are almost equal, and its carotenoid content is higher
compared to that of PS I. the photocentre is a special chlorophyll a molecule called P680,
surrounded by other chlorophylls a and b. PS II also contains Mn2+, Cl-, quencher molecule Q,
plastoquinone, cytochrome complex and plastocyanin. It picks up electron released during
photolysis of water, which is extruded on the absorption of light, and as the extruded electron
passes over the cytochrome complex, sufficient energy is released to take part in the synthesis of
ATP from ADP and inorganic phosphate (Pi) in a process called non-cyclic photophosphorylation.
PSII can only operate in conjunction with PSI.

Photon

Carotenes Carotenes
Chl - b Chl - b
Antenna
Various Molecules Various
forms of forms of
Chl-a Chl-a

P 700 Reaction Centre P 680 Reaction Centre

PS I PS II
Red Drop Effect

Wavelengths beyond 700nm are apparently of insufficient energy to drive any part of
photosynthesis. So a huge drop in efficiency has been noticed at 700nm. This phenomenon is called
as RED DROP EFFECT. In other words there is a sharp decrease in quantum yield at wavelengths
greater than 680nm. This decrease in quantum yield takes place in the red part of the spectrum.
The number of oxygen molecules released per light quanta absorbed is called as quantum yield of
photosynthesis. This effect was first of all noticed by Robert Emerson of Illinois University. Later
on Emerson and his group observed that if chlorella plants are given the inefficient far red light
and red light of shorter wavelengths in alternate fashion, the quantum yields were greater than
could be expected from adding the rates found when either colour was provided alone. This
synergistic effect or enhancement is known as EEE or Emerson Enhancement Effect. This was the
first good evidence that there are two photo systems; one absorbs far red light and other red light
and both of them must operate to drive photosynthesis most effectively.

Enhancement with
supplementary light

0.10
yield

0.08
Red drop
Photophosphorylation

Photophosphorylation is the formation of ATP from ADP and inorganic phosphate (Pi) with the
help of solar energy. It is of two types; cyclic and non-cyclic.

Cyclic Photophosphorylation: Here, the electron extruded by the reaction centre of PS I (P700),
returns back to the reaction centre after passing over a number of electron carriers. It is performed
by PS I independently. The PS I, after gaining photon of energy extrude electron from its reaction
centre. This electron is picked up by FeS and then passes it over to ferredoxin (Fd), cytochrome
b6, and cytochrome f. From cytochrome f, the electron moves to plastocyanin from where it
becomes available to P700 again.

Non-Cyclic Photophosphorylation: Both PS I and II are involved here. Electron extruded by

P680or PSII does not return to it but is passed on to NADP+ with the help of PS I. PS II after gaining
photon energy extrude electron which is picked up by pheophytin, a non-Mg chlorophyll a, from
which electron moves on to another electron acceptor Q and then to plastoquinone (PQ). The
electron is picked up by cytochrome f which passes it on to plastocyanin (PC). The PC then hands
it over to PS I reaction centre (P700), from where it passes over FeS, Fd and then to NADP+.

Cyclic Phosphorylation Non-Cyclic Phosphorylation

1. Only PS I is involved Both PS I and PS II are involved
simultaneously
2. Only longer wavelength of red light is used Utilizes both long and short wavelength of red
light
3. Electrons move in cyclic manner Electrons move in zig-zag manner (Z-scheme)
4. Photolysis of water does not occur and Water is oxidized and oxygen is liberated
hence no evolution of oxygen
5. For each pair of electron transported, only One NAPDH and two ATP are formed for
one ATP is formed, no NADPH every electron transported.
6. It is not inhibited by DCMU Inhibited by DCMU

Mechanism of Photosynthesis

Photosynthesis is the biological process by which some energy rich carbon containing compounds
are produced from CO2 and H2O by the illumination of chloroplast containing cells resulting in
the liberation of oxygen as a by-product. It is essentially and Oxidation-Reduction (Redox)
reaction by which H+ is transferred from water to carbon dioxide through a carrier substance, the
nature of which is still not perfectly understood. In the process, the volume of carbon dioxide
absorbed is almost equal to that of oxygen liberated. Since glucose or fructose appears to be the
first carbohydrate formed, the overall reaction can be represented as:
 Light
6CO 2 + 6H 2O
Chlorophyll
C6H12O6 + 6H 2O + 6O 2

The above equation does not completely explain the sequence of complex reactions involved. The
complexity is further increased by the fact that in this process, the most important energy
transformation process takes place –the transformation of the absorbed radiant energy into stable,
highly potent chemical energy which is available for all vital activities of the living cell.

Nature of the Reactions in Photosynthesis

Photosynthesis as whole fundamentally consists of redox reaction series in which one compound
(water) is oxidized and another compound (carbon dioxide) is reduced. The reactions in
photosynthesis have been grouped into two phases. This was first reported by Blackman in 1901:

1. Light Reaction – for which light is essential

2. Dark Reaction – which can proceed without light

Light Reaction

This phase of reaction in photosynthesis is also known as Hill’s or Photochemical reaction. It

occurs in the grana (thylakoids) of the chloroplast. The rate of reaction is usually regulated by the
intensity and quality of light and the light capturing pigments. Light reaction can be further divided
into two phases; the photolysis of water leading to the evolution of oxygen and synthesis of
assimilatory power.

Photolysis of Water: It is the breaking up of water into hydrogen and oxygen in the illuminated
chloroplast. A small complex of protein attached to the PS II called oxygen evolving complex
(OEC) oxidizes water into oxygen, H+ and electrons. Cl-, Ca+ and Mn2+ are required during this
process.

Synthesis of Assimilatory Power: Electrons released during photolysis of water are picked up by
the reaction centre of PS II (P680). On receiving a photon of light energy, P680 expels an electron.
The electron extruded by P680 is picked up by pheophytin from where electron passes over a series
of carriers. While passing over cytochrome complex, the electron loses sufficient energy for
creation of proton gradient and synthesis of ATP (photophosphorylation). From plastocyanin, the
electron is picked up by the reaction centre (P700) of PS I. Electron is expelled by P700, it is picked
up by FeS, ferredoxin and NADP reductase. The NADP reductase releases the electron to NADP+
to enable it combine with the H+ produced in the photolysis of water to produce NADPH.

Dark Reaction

As the name implies, this phase of photosynthetic reaction does not require the presence of light.
During this reaction or biosynthetic pathway, assimilatory power is used in the fixation and
reduction of CO2. The enzymes required for this reaction are present in the matrix or stroma of
chloroplast. There are two main pathways; C3 or Calvin Cycle and C4 or Hatch and Slack Cycle.

Calvin or C3 Cycle: this path of carbon assimilation was given by Calvin, Benson and Bassham
(1949). It is called C3 cycle because carbon dioxide reduction is a cyclic process and the first stable
product in the cycle is a 3-carbon compound called 3-phosphoglyceric acid or 3-PGA.
The cycle spends ATP as an energy source and consumes NADPH2 as reducing power for adding
high energy electrons to make the sugar. There are three phases of the cycle. In phase 1 (Carbon
Fixation), CO2 is incorporated into a five-carbon sugar named ribulosebisphosphate (RuBP). The
enzyme which catalyzes this first step is RuBP carboxylase or rubisco. It is the most abundant
protein in chloroplasts and probably the most abundant protein on Earth. The product of the
reaction is a six-carbon intermediate which immediately splits in half to form two molecules of 3-
phosphoglycerate. In phase 2 (Reduction), ATP and NADPH2 from the light reactions are used to
convert 3-phosphoglycerate to glyceraldehyde 3-phosphate, the three-carbon carbohydrate
precursor to glucose and other sugars. In phase 3 (Regeneration), more ATP is used to convert
some of the pool of glyceraldehyde 3-phosphate back to RuBP, the acceptor for CO2, thereby
completing the cycle. For every three molecules of CO2 that enter the cycle, the net output is one
molecule of glyceraldehyde 3-phosphate (G3P). For each G3P synthesized, the cycle spends nine
molecules of ATP and six molecules of NADPH2. The light reactions sustain the Calvin cycle by
regenerating the ATP and NADPH2.

Photorespiration: Under conditions of high O2 and low CO2, rubisco has an oxygenase activity,
initiating the phenomenon called photorespiration. When this happens, 3-phosphoglycerate and
phosphoglycolate are formed in the chloroplast from ribulose-1,5-bisphosphate. The
phosphoglycolate is then dephosphorylated and transferred into the peroxisomes (cell organelles)
where glycolate is oxidized to H2O2 (destroyed by catalase) and glyoxylate. Glyoxylate is aminated
to produce glycine, which is transferred to the mitochondria. There, two glycines are converted
into one molecule of serine plus one molecule each of CO2 and NH3. The serine ultimately is
converted back to 3-phosphoglycerate.
Photorespiration results in a net loss for the cell: Ribulose-1,5-bisphosphate is lost from the Calvin
cycle;O2 is consumed, CO2 is released; only a part of the carbon is returned to the chloroplast;
ATP is expended without apparent benefit.

Plant cells probably undergo photorespiration for a reason, however. Under conditions of high
illumination, CO2 levels around a plant may fall. Under these conditions, reactive oxygen species,
such as superoxide, could be produced, causing damage to the plant. Photorespiration's function
may be to protect against reactive oxygen species under these conditions by consuming oxygen
and releasing CO2.

C4 or Hatch - Slack Cycle: This cycle was first reported by Kortschak, Hartt and Burr in 1965.
They found that 4-carbon compounds, like aspartic acid and malic acid, result from the fixation of
CO2 instead of PGA. Hatch and Slack (1966) later supported this and reported that Oxaloacetic
acid (OAA), a 4-carbon compound, is the first product of carbon fixation. This cycle is found in
important crop species, such as maize and sugarcane, and is important in tropical plants, which are
exposed to intense sunlight and high temperatures. Photorespiration is most active under these
conditions. The C4 cycle efficiently utilizes the CO2 released as a result of photorespiration.
The CO2 acceptor molecule here is the phosphoenol pyruvate (PEP) and not the RuBP.
Furthermore, PEP-carboxylase is the key enzyme in the C4; Rubisco is negligible or completely
absent in the mesophyll chloroplast, but present in the bundle sheath chloroplast.

C4 Plant: the C4 plants, mostly monocots, have a characteristics leaf anatomy called Kranz
Anatomy. Here the vascular bundles are surrounded by sheath of large parenchymatous cells called
bundle sheaths which are surrounded by mesophyll cells. These plants have two types of
chloroplast (chloroplast dimorphism);

- Bundle sheath chloroplast: they are large, lack grana and contain starch grains
- Mesophyll chloroplast: smaller in size, contain grana and lack starch grains.
Differences between the C3 and C4 Pathways

C3 Pathway C4 pathway
1. Usually seen in temperate plants and Common in tropical and sub-tropical crops
some tropical plants
2. Leaves do not show Kranz anatomy Leaves show Kranz anatomy

3. Green cell show only grana chloroplast Mesophyll cells have agrana chloroplast and
bundles sheath cells have grana chloroplast
(chloroplast dimorphism)
4. Only calvin cycle is operated C4 pathway in mesophyll cells and C3 in
bundle sheath cells
5. Primary CO2 acceptor is RuBP Primary CO2 acceptor is PEP

6. The first formed stable compound is PGA The first formed stable compound is OAA

7. Less efficient in CO2 fixing More efficient

8. Show high photorespiration Photorespiration is negligible

9. Optimum temperature is 15 - 25°C 30 - 45°C

10. Photosynthetic yield is low to high High

11. 18 molecules of ATP are used for the 30 molecules of ATP are used; 18 in the
formation of one molecule of glucose bundle sheath cells and 12 in the mesophyll
cells
12. Less efficient in water usage Water usage efficiency is high

13. CO2 compensation point is very high Low

Crassulacean Acid Metabolism (CAM) Cycle: This metabolism was first reported in
Bryophyllum, a member of Crassulaceae family, hence the name CAM. It occurs mostly in
succulents (xerophytes) like Agave, Aloe, Pineapple, etc. These plants have scotoactive stomata,
which are open at night and closed during most part of the day. They demonstrate what is known
as dark acidification. During the night, malic acid is formed by utilizing the CO2 that diffuses
through the open stomata. The malic acid formed is broken down into CO2 and pyruvic acid in the
day time, the CO2 is then released for utilization in the Calvin cycle. Two main enzymes are
involved; PEP carboxylase during the night and Rubisco during the day.

Blackman’s Law of Limiting Factor

Blackman (1905) stated that “when a process is conditioned as to its rapidity by a number of
separate factors, the rate of the process is limited by the pace of the slowest factor”.

Light intensity
G H
High
Rate of Photosynthesis

E F Medium

C D Low

A CO2 Concentration B

Factors Affecting Photosynthesis

Light intensity, temperature, carbon dioxide and water are the chief external conditions for
photosynthesis and its rate. The chief internal factors are chlorophyll content, accumulation of
photosynthetic products and protoplasmic factor.

1. Light: No photosynthesis occurs in a very weak light. As the intensity of light increases, the
process begins and picks up the pace. A very strong intensity of light causes solarization.
During solarization, photo-oxidation occurs and if it continues for a few hours, the
photosynthetic apparatus is destroyed. Photosynthesis begins in the morning hour, reaches its
 peak in the noon/afternoon and then its rate declines. The value of light at which further
increase is not accompanied by an increase in CO2 uptake is called light saturation point. At
this point, a plant continues to respire almost at a uniform rate. During peak hours, rate of
photosynthesis is about 20 times faster than that of respiration. Light compensation point is
however reached in the morning as well as in the evening.
2. Temperature: Plants can perform photosynthesis on a range of temperatures. While some
cryophytes can photosynthesize at -35°C, some thermal algae can photosynthesize even at
75°C. Usually, plants can perform photosynthesis at between 10 - 40°C. The optimum
temperature ranges between 25 - 30°C.
3. Carbon Dioxide: the concentration of CO2 in the atmosphere is 0.03% by volume. If this
concentration is increased by 15 to 20 times, the photosynthetic rate increases, if no other
factor becomes limiting. Higher concentration of CO2 can become toxic to plant. However,
the value of the upper limit of CO2 concentration is variable. A stage in CO2 concentration
where there is no net absorption of CO2 by illuminated plant organ is called CO2
compensation point or threshold value. It is less than 10 ppm in C4 plants and ranges
between 50 – 100 ppm for C3 plants.
4. Water: A plant utilizes less than 1% water from its total absorption and the rest is transpired.
A decrease in water contents brings about loss of turgor leading to the closure of stomata. The
effect of water is more of indirect than direct.
5. Oxygen: The concentration of oxygen in the atmosphere is about 21% by volume and it
seldom fluctuates. An increase in oxygen concentration brings about a reduction in
photosynthesis and the phenomenon is called Warburg effect. In C3 plants, the rate of day
respiration is faster than the dark respiration because of photorespiration which occur in high
light intensity, high oxygen conc. And high temperature. Here, oxygen competes with CO2 for
the oxidation of RuBP to PGA thus reducing the fixation of CO2.
6. Chemicals: Chemicals which act as enzyme inhibitor like all other vital processes, inhibit
photosynthesis also. Such chemicals are cyanides, hydroxyl amine, H2S, CO, indoacetate, etc.
Besides, chloroform and ethers also inhibit photosynthesis. The herbicides dichlorophenyl
dimethyl urea (DCMU) and chlorophenyl dimethyl urea (CMU) are also photosynthetic
inhibitors. They inactivate PS II thus inhibiting Hill’s reaction.
7. Chlorophyll: This is essential for photosynthesis. The plastids are powerless without
chlorophyll. The same reason why the non-green parts of plant cannot photosynthesize.
8. Accumulation of Photosynthetic Products: A heavy accumulation of these products,
whether sugar or starch , in the assimilating cells causes the process of photosynthesis to slow
down and even come to a standstill.
9. Protoplasmic Factor: An internal factor other than chlorophyll plays a part in photosynthesis.
This is called protoplasmic factor. It may be enzymatic in nature.
10. Minerals: Mn2+, Cl-, Ca2+ ions help in photolysis of water. Mg and Fe in synthesis of
chlorophyll. B and K help in translocation of solutes.
11. Hormones: Auxin, Gibberellin and Cytokinin increase photosynthesis. ABA decreases it.

Photosynthesis in Bacteria

There are certain groups of bacteria with pigments. These are photosynthetic autotrophic bacteria.
Their pigments are closely related to the chlorophyll found in higher plants. They contain three
types of pigment, namely bacteriovirdin, bacteriochlorophyll and carotenoids. There are three
groups of photosynthetic bacteria, they include:

1. Green sulphur bacteria: these contain the bacteriovirdin green pigment. They carry out
photosynthesis by utilizing H2S instead of H2O as hydrogen donor. Examples are
Chlorobium sp. and Chlorobacterium sp.

6CO2 + 12H2S C6H12O6 + 6H2O + 12S

2. Purple bacteria: these can either be purple surphur or non-sulphur. The purple sulphur
bacteria contain the reddish purple pigment called bacteriochlorophyll which is similar to
chlorophyll a. Here, the hydrogen donors are sulphur compounds and molecular hydrogen.
It absorbs invisible infra-red light of wavelength 800 – 890 nm. An example is the
Chromatium sp. The purple non-sulphur bacteria also contain bacteriochlorophyll but here
photosynthesis takes place in the presence of simple organic compounds such as acids and
alcohols. An example is Rhodospirillumrubrum.
3. Heliobacteria: these are recently (1983) discovered photosynthetic bacteria which contain
a new type of bacteriochlorophyll called bacteriochlorophyll-g as antenna and reaction
centre pigment. They also utilize organic compounds as electron donors.
 RESPIRATION

Respiration is a controlled cellular process of oxidation and decomposition of organic compounds,

particularly simple carbohydrates such as glucose, leading to the release of energy. It is thus a
catabolic process. The potential energy stored in the organic compounds in living cells is by this
oxidative process released in a stepwise manner in the form of kinetic energy under the influence
of a series of enzymes, and this energy released is made available to the protoplasm for its manifold
vital activities such food manufacture, movement and growth. A considerable amount of energy
however often escapes in the form of heat as can be seen in germinating seeds.

The reserve food materials that undergo oxidation are mostly simple carbohydrates, principally
glucose, and sometimes also, particularly in the absence of glucose, other substances such as
complex carbohydrates, proteins and fats. These complex substances are first hydrolyzed and then
oxidized to release energy.

The main facts associated with respiration are:

- Consumption of atmospheric oxygen

- Oxidation and decomposition of a portion of stored food, resulting in loss of dry weight
usually observed in seeds germinating in the dark
- Liberation of carbon dioxide and a small quantity of water (the volume of CO2 liberated
being equal to the volume of O2 consumed)
- And most importantly, the release of energy by the breakdown of organic food.

The overall reaction can be given as follows:

C6H12O6 + 6O 2 6CO 2 + 6H 2O + Energy (2868 KJ)

In the above reaction, oxygen serves as the electron acceptor leading to the formation of water,
and carbohydrate is oxidized to CO2.

The oxidation process in repiration may be complete as cen be seen in the above reaction with the
formation of CO2 and H2O as the end product. This is aerobic respiration. CO2 escapes from the
plant body while H2O gets mixed up with the general mass of water in the cell. The oxidation
process may also be incomplete leading to the formation of some organic acid or ethyl alcohol as
shown below:

C6H12O6 2C 2H5OH + 2CO 2 + Energy (118 KJ)

This is anaerobic respiration.

Aerobic and Anaerobic Respiration

Normally, free oxygen is used in respiration, resulting in complete oxidation of stored food and
formation of carbon dioxide and water as end-products. This is known as aerobic respiration. A
considerable amount of energy is released by this process. Under certain conditions, especially in
the absence of free oxygen, many tissues of higher plants, seeds in storage, fleshy fruits and
succulent plants take to a kind of respiration called anaerobic respiration. Such respiration results
in incomplete oxidation of stored food and formation of carbon dioxide and ethyl alcohol, and
sometimes also various organic acids such as malic, citric, oxalic, tartaric, etc. The energy released
by this process is not sufficient to maintain the activity of the protoplasm.

Mechanism of Aerobic Respiration

Chemical changes in aerobic respiratory process occur in three distinct phases and is controlled by
a group of different kinds of complex enzymes which work step by step. The first phase, which is
the anaerobic phase, is the incomplete oxidation of glucose to pyruvic acid through a chain of
intermediate reactions called Glycolysis. This phase does not require the use of free oxygen. The
second phase involves the complete oxidation of pyruvic acid to CO2 and H2O in a cycle of
reactions called Krebs Cycle, with each reaction being controlled by specific enzyme. It takes
place in the presence of oxygen. The third phase is called the Electron Transport System (ETS).

Anaerobic Phase (Glycolysis)

In the first phase of respiration, simple carbohydrate like glucose or its isomer, fructose, is first
phosphorylated (addition of a phosphate group by ATP). In this process, ATP is converted to ADP.
This is the initiating step of the respiratory process leading to the formation of phosphoglyceric
acid and, finally, pyruvic acid.The scheme of glycolysis was given by Gustav Embden, Otto
Meyerhof and J Parnas. Due to this, it is also called the EMP Pathway.
The reactions take place in the cytoplasm of the cell. The overall reaction is:

Glucose+2ATP+4ADP+2NAD→2 Pyruvic acid+2ADP+4ATP+2NADH

The following points may be noted:

1. 1 mol. of glucose yields 2 mol. of pyruvic acid

2. The phosphate group as a source of energy is donated mainly by ATP
3. Each reaction is catalyzed by a specific enzyme
4. All the reactions are reversible
5. Pyruvic acid holds the intermediate key position between the two phases of respiration.

Anaerobic Oxidation of Pyruvic Acid

In the absence of oxygen, pyruvic acid is converted to acetaldehyde and CO2 under the action of
pyruvic decarboxylase. Acetaldehyde is now the starting point from which reactions leads to the
production of a number of compounds. In anaerobic bacteria and some fungi, the absence of
oxygen causes the conversion of pyruvic acid to certain compounds under the action of certain
enzymes with the release of certain amount of energy which is however too small for the activity
of such microorganisms. Thus in some bacteria, pyruvic acid is reduced to lactic acid while in
others it is butyric acid. The fermentation of glucose or anaerobic respiration in yeast cells leads
to the conversion of pyruvic acid to ethyl alcohol and CO2. In higher plants, anaerobic respiration
may continue only for a short period because the end-products formed in the process are mostly
toxic.

Aerobic Phase

This phase immediately follows the first phase with the supply of oxygen, and most of the pyruvic
acid is completely oxidized to CO2 and H2O with the liberation of a maximum amount of energy.
The ATP formed is about four times greater than that formed in glycolysis. Several organic acids
and amino acids are formed during the process of aerobic respiration. A series of reactions takes
place in a step-wise manner in the whole process under the action of distinct complex enzymes.

The sequence of reaction in this phase may be divided into two:

1. Formation of Acetyl-CoA, and

2. Citric Acid Cycle or Tri-Carboxylic Acid Cycle or Krebs’ Cycle

Formation of Acetyl-CoA: With the access of sufficient oxygen, the pyruvic acid formed in
glycolysis undergoes oxidative decarboxylation and is converted to Acetyl-Coenzyme A in the
matrix of mitochondria. The chemical process leading to this conversion is however very complex,
involving a series of reactions. Briefly speaking, pyruvic acid reacts with co-enzyme A and NAD+
and is oxidativelydecarboxylated into Acetyl-CoA molecule in the presence of pyruvate
dehydrogenase complex. One mol. of CO2 is released and NAD+ is reduced to NADH + H+. The
reaction is irreversible.

Pyruvic acid + CoA + NAD Acetyl CoA + CO2 + NADH

Citric Acid Cycle or Tri-Carboxylic Acid Cycle or Krebs’ Cycle:The cycle was discovered by
Hans Krebs in 1937 in certain nematodes and pigeon muscles for which he received Nobel Prize
in 1953. The cycle occurs only under aerobic conditions as it requires the continuous supply of
NAD and FAD. In eukaryotes, the reactions in the Krebs cycle occur in the matrix of mitochondria
while in prokaryotes, it occurs in the cytoplasm. Pyruvic acid enters the matrix of mitochondria
where it undergoes chemical reaction producing 3 molecules of CO2 and NADH. At the end, the
Krebs’ cycle starting compound, a 4C Oxalo Acetic Acid (OAA), is regenerated. The cycle is
initiated with the formation of Acetyl CoA from pyruvic acid. Acetyl CoA condenses with a 4C
compound called Oxalo acetic acid (OAA), forming a 6C citric acid. As the first compound formed
during kreb’s cycle is citric acid, this cycle is also called ‘citric acid’ cycle. Organic acids
containing three carboxylic acid groups (COOH) are also formed during this cycle. Hence it is also
called ‘tricarboxylicacid’cycle. The overall reaction of citric acid cycle is,

Pyruvic acid+4NAD+FAD+2H2O+ADP+Pi → 3CO2↑+4NADH+4H++FADH+ATP

Carbon dioxide is released to the atmosphere or green cells may use it for photosynthesis. The
hydrogen which is present in glucose will be used for reducing NAD and FAD to NADH and
FADH2 respectively.

Electron Transport System/Oxidative Phosphorylation: It is third step of aerobic respiration,

taking place in the inner mitochondrial membrane (Cristae). In this step of respiratory process,
NADH and FADH2 formed during glycolysis and Kreb’s cycle are oxidized to NAD and FAD.
During this oxidation process, electrons pass through series of electron carriers (like co-enzyme
Q, cytochromes b, c, a and a3) and ADP is phosphorylated to ATP by the addition of Pi. Since
oxidation of NADH and FADH2 is associated with the synthesis of ATP, it is called Oxidative
Phosphorylation. Oxidation of one NADH and one FADH2 yields three andtwo ATP molecules
respectively. The hydrogen atom (H+) and the electron (e-) of the reduced NADH and FADH2 are
accepted by the oxygen to form a molecule of water and it is called terminal oxidation. Total gain
of ATP molecules at the end of complete oxidation of 1 molecule of glucose is 38.

Amphibolic Pathway
During aerobic respiration the organic substrates are broken down into simple substances such as
carbon dioxide and water (Catabolism). However it also produces many intermediate organic
compounds/acids during kreb’s cycle which form the starting compound for the synthesis of other
complex substances, needed for the cell. Hence respiration is said to be both catabolic and anabolic
and is referred to as ‘amphibolic pathway’.

Respiratory Quotient
Respiratory quotient is defined as the ratio of volume of CO2 evolved to the volume of Oxygen
consumed during respiration. The volume of carbon dioxide released and the volume of oxygen
consumed by the cell during respiration depends upon the type of respiratory substrate utilized.
Therefore RQ value indicates the substrate type used for respiration.
𝑉𝑜𝑙𝑢𝑚𝑒 𝑜𝑓 𝐶𝑂2 𝐸𝑣𝑜𝑙𝑣𝑒𝑑
𝑅𝑄 =
𝑉𝑜𝑙𝑢𝑚𝑒 𝑜𝑓 𝑂2 𝐶𝑜𝑛𝑠𝑢𝑚𝑒𝑑
The RQ for carbohydrate aerobic respiration is 1. That is the rate of CO2 evolve is equal to the rate
of O2 consumed. RQ of protein aerobic respiration is between 0.7 and 0.9 while that of fats and oil
is also less than 1. The value is greater than 1 when the substrate is organic acid. The RQ in CAM
plants such as Bryophyllum is 0 because there is no evolution of CO2 since oxidation occurs in the
dark. It is infinity in anaerobic respiration since there is no consumption of oxygen.

Factors Affecting Respiration

1. Temperature: the optimum temperature for respiration is about 30°C to 35℃, with
minimum being 0°C and maximum 45℃. At low temperatures, the enzymes become inactive
and so the rate of respiration drops. At very high temperatures, respiration also slows down
and may even stop due to denaturation of the enzymes.
2. Oxygen: Complete absence of oxygen results in anaerobic respiration and aerobic
respiration stops. And in aerobic organisms, continued absence of oxygen will result in
death. When sufficient amount of oxygen becomes available, aerobic respiration starts and
anaerobic will be completely stopped. This is known as the oxygen extinction point.
3. Carbon Dioxide: Higher concentration of CO2 in the atmosphere has a retarding effect on
respiration.
4. Inorganic Salts: It has been observed that if a plant is transferred from water to a salt
solution, the rate of respiration increases.
5. Water: Proper hydration of respiring cells is essential for respiration. Rate of respiration
decreases with decrease in the amount of water, so much so that in dry seeds the rate of
respiration is at its minimum. This is because water provides the appropriate medium for the
respiratory enzymes.
6. Light: Light has an indirect effect on the rate of respiration, through its role in the synthesis
of organic matter.
7. Stimulation: Mechanical disturbance, injury or infection increases respiration. Hopkins
(1927) found that in wounded plant tissue the sugar content is suddenly increased which is
responsible for temporary increase in the respiration rate.
8. Respiratory Substrate:Higher availability of respiratory substrates increases the rate of
respiration upto a certain limit.
9. Protoplasmic Factor: Young growing cells exhibit high rate of respiration as compared to
mature cells.
10. Inhibitors:A number of chemicals inhibit respiration e.g., azide, cyanide, malonate, carbon
monoxide etc.

What are the differences between Photosynthesis and Respiration?

Respiration Photosynthesis
Catabolic Anabolic
Raw material = CHO, protein, fat, organic Raw materials = CO2, H2O, chlorophyll, light
acids and oxygen
Occur in all living cells Occur in only green cells
Happens day and night Happens only by day
Exothermic process Endothermic process
End-product = CO2& H2O, or alcohol/malic End-product = CHO and O2
acid
Includes dehydrogenation and decarboxylation Includes hydrogenation and carboxylation
Either completed in cytosol only or cytosol and Completed in chloroplast (eukaryotes) or
mitochondria, or chloroplast, peroxisome, and chromatophores (prokaryotes)
mitochondria
Oxidation of 1 mol of glucose yields 38 ATP 1 mol of glucose synthesized at the cost of 18
mol. ATP mol.
 NITROGEN USE IN PLANTS

Nitrogen is a vitally important plant nutrient. It is now known to be an essential component of

living matter. In plants, nitrogen is utilized for the synthesis of amino acids, which are the building
blocks for proteins. The protein in the vegetative cells of plants is largely functional than structural
in nature. Many of these proteins are enzymes while others are nucleoproteins, some of which are
present in the chromosomes. Proteins thus serve as catalyst and as directors of metabolism.

Apart from its role in the formation of protein, nitrogen is an integral part of the chlorophyll
molecule. The chlorophyll molecule is made up of a central magnesium atom surrounded by 4
pyrol rings, each of which contains one nitrogen and 4 carbon atoms. An adequate supply of
nitrogen to plants has been associated with vigorous vegetative growth and deep green colour
because of its role in chlorophyll formation. Excessive quantities of nitrogen can under certain
conditions prolong and delay crop maturity. This is most likely to occur in the absence of adequate
supply of other nutrients.

The supply of nitrogen is related to carbohydrate utilization. When nitrogen supply is insufficient,
carbohydrate may be deposited in vegetative cell which will cause then to thicken. When nitrogen
supply is adequate and conditions are favourable for growth, proteins are formed from the
manufactured carbohydrates. Less carbohydrate is thus deposited in the vegetative portion, more
protoplasm is formed, and because protoplasm is highly hydrated, a more succulent plant results.
Excessive succulence in some crop may have harmful effect. With crop such as cotton, a
weakening of the fibre may result, and with grain crops, logging may occur, particularly when
potassium supply is inadequate. In some cases, excessive succulence may make the plant become
susceptible to diseases and insect attack.

When used in conjunction with other plant nutrient, in a good crop management programme,
nitrogen greatly increases crop yield. Deficiency in nitrogen results in stunted growth and
yellowing of leaves. This yellowing or chlorosis usually appears first on the lower leaves while
the upper leaves remain green. In case of severe shortage in nitrogen supply, the leaf will turn
brown and eventually die.
SOURCES OF NITROGEN

The chief sources of nitrogen are atmosphere and soil. It exists in molecular (N2) form in the
atmosphere, while in the soil it may exist in inorganic or organic forms. Inorganic forms of nitrogen
in the soil are ammonical nitrogen, nitrate nitrogen and nitrite nitrogen.

Molecular Nitrogen: this is the free nitrogen found in the air; an inert gas which constitue about
76% of air by weight and 78% by volume. This form of nitrogen is not available to plant directly.
Unlike oxygen and carbon dioxide, nitrogen cannot diffuse inside or outside of the cell through
stomata.

Ammonical Nitrogen: this is nitrogen in the form of ammonia. Ammonia is formed by fixation
of free atmospheric nitrogen by physical (lightening), chemical (industrial) and biological means.
It is also formed from organic compounds by several types of micro-organisms. This form of
nitrogen get mixed into the soil and absorbed through the roots.

Nitrate Nitrogen:Ammonical nitrogen gets oxidized by Nitrosomonas bacteria to nitrites and then
to nitrates by Nitrobacter bacteria. Nitrates are also produced by physical means. This forms of
nitrogen is readily available for plant growth.

Nitrite Nitrogen: this form of nitrogen is also absorbed by plants and also formed by reduction of
nitrate.

Organic Nitrogen: this includes amino acids, urea and amides. They are formed in the soil by
degradation of dead remains of animals and plants.

NITROGEN FIXATION

Nitrogen fixation refers to the oxidation or reduction of atmospheric nitrogen to a form that can be
utilized by living organism. The usable forms are most frequently nitrate or ammonia, but some
small organic molecules such as urea and amino acids can be taken up. In nature, nitrogen can be
fixed in three ways:

- Photochemical Reactions. The gaseous nitric acid (NO) and ozone (O3) react together in
the presence of light to produce nitric acid (HNO3). This accounts for about 2% of the
nitrogen fixed. All of the organisms
- N2 Fixation by Lightning. Lightning causes the formation of highly reactive OH- radicals,
H and O from water vapour (H2O) and oxygen (O2). These products react with nitrogen to
form nitric acid which comes down in rain water. Lightning accounts for about 8% of the
total nitrogen fixed on earth
- Biological Nitrogen Fixation. This is brought about by either non-symbiotic or by
symbiotic association with several bacteria or cyanobacteria (blue-green algae) which are
provided by nif gene and nitrogenase enzyme. This account for the remaining 80% of the
nitrogen fixed on earth.

Biological Nitrogen Fixation

The organisms capable of fixing atmospheric nitrogen are prokaryotes which can either be
symbiotic or non-symbiotic (free living).

Symbiotic N-fixing Bacteria: Bacteria of the genus Rhizobium are known to live in association
with legumes. The infection process forming the symbiosis has been studied in much detail. During
the actual infection process that must involve the production of hormones and digestive enzymes,
the root hairs form a coil and then is invaded by the bacteria. Once in the root hairs, migration to
the root cortex takes place via a mucilaginous infection thread. After the bacteria enter the cortical
cells, they cause it to increase in size and ultimately form the nodules on the upper surface of the
root. After their release into the cortical cells, the rhizobia stop dividing, and they enlarge. The
enlarged, non-motile groups of bacteria inside the membranes of the host cells are called
bacteroids.

Although the legume nodule containing N-fixing bacteria Rhizobium spp. is the most well-known,
there are other non-legumes that have nodules with N-fixing organism. Many of these nodules
occurring with non-legume have actinomycetes of the genus Frankia as the N-fixing organism.
More recently, it has been shown that certain tropical grasses including paspallum, digiteria, maize
and sorghum have symbiotic non-nodule forming bacteria of the genera azotobacter, spirillum and
klebsiella that fix nitrogen. These bacteria appear to live in the mucilaginous plate along roots. To
date, estimate of the capacity for nitrogen fixation by these organisms which live in association
with grasses have been so small compared to the rhizobia legumes.
Non-Symbiotic N-Fixation:As a result of the extremely high energy demand of the nitrogen
fixation process, many of the nitrogen fixing organisms are symbionts. Nevertheless, there are
varieties of non-symbiotic organisms capable of nitrogen fixation. The non-symbiotic nitrogen
fixing bacteria include free living heterotrophic bacteria, photosynthetic bacteria and blue-green
algae. The heterotrophic bacteria obtain the required energy from the environment, but most
certainly, the energy for nitrogen fixation by the photosynthetic organisms comes chiefly from
photosynthetic process.

The free living heterotrophic nitrogen fixing bacteria fall into three groups:

- Obligate Aerobes
- Facultative Aerobes
- Obligate Anaerobes

The obligate aerobes include the species of the azotobacteriaceae and some members of the
pseudomonaceae. The facultative aerobes include species of bacteria such as Klebsiella and
Bacillus. Examples of anaerobic nitrogen fixing organisms include Clostridium and
Methanococcus.

The nitrogen fixing photosynthetic bacteria are gram negative bacteria living in aquatic
environment. There is much evidence that these photosynthetic organisms are directly involved in
nitrogen fixation process. Many species of the blue-green algae have the capacity of nitrogen
fixation. The nitrogen fixing blue-green algae are mostly of the types that have heterocyst.
Evidently because oxygen evolved by the green portion of the algae inhibits nitrogen fixing
enzymes, the heterocyst may be a specialized structure in which there is reduced oxygen tension
during nitrogen fixation. It should be noted that the photosynthetic bacteria do not evolve oxygen
during photosynthesis.

The Biochemistry of Nitrogen Fixation

The produce of nitrogen fixation by intact cells is ammonia. Cell-free preparations and purified
protein have shown that an enzyme called nitrogenase catalyzes the reduction of nitrogen to
ammonia. The enzyme has four basic requirement for activities in vivo. There is a strong
requirement for (i) ATP, (ii) magnesium cation, (iii) a reducing agent as a source of electron for
the reduction, and (iv) a need for anaerobic condition because nitrogenase is sensitive to oxygen.
The reaction as presently understood is as follows:

N2 + 6e-+ 8H
+ +
+ 2Mg + 2ATP 2NH 3 + H2 + 2MgADP + 2Pi

The multi-step reduction process is believed to proceed from molecular nitrogen to diamide to
hydrazone and ultimately to ammonia.

NH3 HN NH NH2 NH2 2NH3

Diamide Hydrozone Ammonia

Each of the above steps require two electrons, making a total of six to reduce one diatomic nitrogen
molecule to two ammonia molecules. The source of electron for the reaction is ferredoxin
especially in the case of photosynthetic bacteria and blue-green algae capable of nitrogen fixation.
Ferredoxin is reduced directly in photosynthesis and then used as reductant in nitrogen fixation.

2 MgADP + 2Pi

Fe Protein MoFe Protein

Ferredoxin N2 + 8H+
(Oxidised) (Reduced)
(Reduced)

Fe Protein MoFe Protein

Ferredoxin 2NH3 + H2
(Reduced) (Oxidised)
(Oxidised)

2 MgATP

Nitrogenase enzyme complex

PLANT GROWTH AND DEVELOPMENT

The growth and development of flowering plant is a complex phenomenon. It can be considered
to begin with germination followed by a large complex of series of physiological and
morphological events that are called growth and development. After germination and vegetative
growth, the vegetative apexes will differentiate to form reproductive apexes. The reproductive
apexes produce flowers or fruiting structures, and ultimately fruits and seeds. The seed that is
produced goes into a dormant stage prior to germination which will complete the cycle.

In the case of annual plants, the vegetative portion will die and only the seeds carry on to the
following growing season. Perennial plants do not die but frequently go into a dormant stage prior
to resuming vegetative growth. Throughout the life cycle of a plant, many events that govern
growth and development take place.

The plant is in part a consequence of genetic makeup and in part the consequence of the
environment in which it grows and develops. The development of the plant is regulated internally
by hormones and the entire sequence of the event is modified by the environment. The major
classes of plant hormones, also called phytohormones, are Auxin, Gibberellin, Cytokinins,
Abscisic acid (ABA) and Ethylene.

Specifically, the best known plants’ responses to these various hormones are:

i. cell enlargement for auxin

ii. stem elongation for gibberellin

iii. Cell division for Cytokinins

iv. Dormancy or arrested growth for abscisic acid, and
v. Maturation for ethylene.

Definition of Growth and Development

Growth is an extremely complex natural phenomenon and as a result, it’s quite difficult to define
precisely. Growth has been simply defined as an irreversible increase in size or volume
accompanied by the biosynthesis of new protoplasmic constituents. Development can then be said
to be a combination of both growth and cellular differentiation. When defined in such a manner,
growth is the quantitative aspect of development representing an increase in the number and size
of cells. Differentiation is the qualitative aspect of development.

Growth can be divided into two processes:

a. Cell division, and

b. Cell enlargement or elongation.

Cell division results in two daughter cells arising from a mother cell, increasing the number of
cells in an individual. It is considered complete when the sizes of the daughter cells have reached
the size of the mother cell.

Cell enlargement or elongation results in an increase in the size of the newly formed cells beyond
that of the mother cell.

Phases of Growth

Growth does not take place throughout the length of the plant body but it is localized in special
regions called meristems, which may be apical, lateral or intercalary. The growth in length is due
to gradual enlargement and elongation of the cells of the apical meristems, located at the root and
shoot apices.

In dicotyledons and gymnosperms, the growth in thickness is due to the activities of the lateral
meristems, i.e. intervascular cambium, vascular cambium and cork cambium. If the history of
growth of any organ of a plant is followed, three phases can be recognized:

1. Formative Phase: this is restricted to the apical meristems. The cells of these regions are
constantly dividing and multiplying in number. These cells are characterized by abundant
protoplasm, a large nucleus and a thin cellulose wall.
2. Elongation Phase: this lies immediately behind the formative phase. The cells no longer
divide but increase in size; they begin to enlarge or elongate until they reach their maximum
size.
3. Maturation Phase: this phase lies further back. Here the cells haven already reached their
permanent size, the thickening of the cell walls begins to take place. This phase is also
known as cell differentiation phase.
Conditions Necessary for Growth

Since growth is brought about by the protoplasm, the conditions necessary for growth as those that
maintain the activities of the protoplasm and they are as follows:

1. Supply of Nutritive Materials: growth can only take place when the protoplasm of the
growing region is supplied with nutritive materials. The protoplasm assimilates these
materials and builds up the body of the plant. Food materials are also a source of energy
for the actively dividing cells.
2. Supply of Water: an adequate supply of water is absolutely necessary to maintain the
turgidity of the growing cells. Turgidity is the first step towards growth. The protoplasm
can only work when it is saturated with water. An abundant supply of water will make up
for loses due to transpiration. It is a fact however that only a small quantity is required for
actual growth.
3. Supply of Oxygen: supply of free oxygen is indispensible for the respiration of all living
cells. Respiration is an oxidative process by which the potential energy stored in food is
released in the form of kinetic energy and made use of by the protoplasm for its manifold
activities.
4. Suitable Temperature: the protoplasm requires a suitable temperature for its activities.
At low temperatures, it stops to perform its functions or does so very slowly while the
temperature of between 45 - 50℃ coagulates and kills the protoplasm. The protoplasm
manifests its activities within a certain range of temperatures; the optimum ordinarily
ranges from 28 – 30℃ and the minimum lies at about 4℃.
5. Light: light is not absolutely necessary for the initial stages of growth. In fact, plants grow
more rapidly in the dark than in the light. The protoplasm maintain its healthy condition
resulting in sturdy plant with normal stem and green leaves with the supply of certain
intensity of light. However, as have been learnt already, light is an essential factor in the
food manufacturing process of plant. The continued absence of light is very harmful to
plant. Plants grown in the dark or in very weak light normally have delicate soft and slender
stems with elongated internodes. They are pale green or pale yellow in colour and sickly
in appearance, they seldom produce flowers and fruits. The leaves are usually small and
often remain underdeveloped. The roots are also seen to be poorly developed. Such plants
are said to be etiolated.
 6. Force of Gravity: this factor determines the direction of growth of particular organs of the
plant body. The root responds positively to force of gravity in its growth while the stem
grows against the force of gravity.

Measurement of Growth

Frequently, an estimate of the growth of a plant can be obtained with a simple ruler or balance.
However, with our definition of growth as an irreversible increase in volume or size, growth
determination must be an indication of irreversible increase and not just simple turgor changes that
are as a result of varying water content. Despite such problems, growth is commonly estimated
from measurement of length, area or weight.

i. Linear Measurement: linear measurement of stem growth is ordinarily a good indication

of the growth of the entire plant since growth is highly coordinated. Linear measurements are also
useful if non-destructive sampling is necessary. A viable technique is to obtain a correlation
between a volume or weight and a linear measurement, allowing a prediction of volume or weight
from subsequent linear measurement.

The following equation will meet most experimental needs:

log 𝑦 = 𝑘 log 𝑥 + log 𝑏

𝑦 = 𝑣𝑜𝑙𝑢𝑚𝑒 𝑜𝑟 𝑤𝑒𝑖𝑔ℎ𝑡

𝑥 = 𝑙𝑖𝑛𝑒𝑎𝑟 𝑚𝑒𝑎𝑠𝑢𝑟𝑒𝑚𝑒𝑛𝑡

𝑘 = 𝑐𝑜𝑟𝑟𝑒𝑙𝑎𝑡𝑖𝑜𝑛 𝑐𝑜𝑒𝑓𝑓𝑖𝑐𝑖𝑒𝑛𝑡

𝑏 = 𝑐𝑜𝑛𝑠𝑡𝑎𝑛𝑡

ii. Leaf Area: In many kind of growth analysis, leaf area is a useful indication. Leaf area can
be determined using a specially designed instrument that makes use of light absorption to measure
the area, or by photographic technique in which a leaf is placed against light sensitive paper. Non-
destructive estimate of leaf area can be obtained by ascertaining the correlation between leaf area
and the linear measurement of the leaf’s length and width. The function that most frequently fits
experimental data is:
𝐴 = 𝑘𝐿 × 𝑊

𝐴 = 𝑎𝑟𝑒𝑎 𝑜𝑓 𝑡ℎ𝑒 𝑙𝑒𝑎𝑓

𝐿 = 𝑙𝑒𝑛𝑔𝑡ℎ 𝑜𝑓 𝑙𝑒𝑎𝑓

𝑊 = 𝑤𝑖𝑑𝑡ℎ 𝑜𝑓 𝑙𝑒𝑎𝑓

𝑘 = 𝑐𝑜𝑟𝑟𝑒𝑙𝑎𝑡𝑖𝑜𝑛 𝑐𝑜𝑒𝑓𝑓𝑖𝑐𝑖𝑒𝑛𝑡

iii. Net Productivity: In growth study in which the objective is to determine total increase in
dry matter (i.e net productivity), a rather rigorous definition has been established by the
International Biological Programme (IBP). The definition of primary productivity is:

𝑃𝑛 = ∆𝐵 + 𝐿

𝑃𝑛 =net dry matter production during a specified time increment between two
measurements

∆𝐵 =total dry weight increase during specified time increment

𝐿 =loss in dry matter during the time increment because of death or consumption by
herbivores.

Net dry matter production does not include losses due to respiration. If losses by respiration is
included, the estimate is gross productivity (Pg). Net productivity is a useful index of growth in
physiological study.

Growth Curves

Grand period of
Growth

growth
Lag phase

Time
A typical growth curve normally shows an increase in size against time. The characteristic for
virtually all growth is a lag phase at the beginning in which growth with time is slow but still
occurs at an increasing rate. A logarithmic phase in which the growth rate is proportional to time
follows the lag phase, and it is called the grand period of growth. After this phase, a decreasing
growth phase occurs, and then finally, a stationary growth phase occurs in which there is no longer
any increase in size with time. The sigmoid (S) shaped growth curve is characteristic of virtually
all growth in living organisms.

Plastochron Index (PI)

Although many indices of plant growth are being developed, most give an estimate of
chronological development. However, in many kinds of studies, analysis according to
chronological age is not of sufficient value because different plant will reach the same
developmental age at different chronological ages/size. It was for this reason that plastochron
index was developed.

PI is a way of measuring the age of a plant dependent on morphological traits rather than on
chronological age. Use of these indices removes differences caused by germination, developmental
differences and exponential growth.

The PI or plastochron age for a plant is the number of leaves that have been developed. For
instance, the plastochron age will be six if the number of leaves developed is six. If exactly five
leaves are developed, the plastochron age will be five. Assuming that the time period during
development of successive leaf is constant, the

log 𝑙𝑛 − log 10
𝑃𝐼 = 𝑛 +
log 𝑙𝑛 − log 𝑙𝑛 + 1

𝑛 =serial number of the leaf that comes closest to exceeding 10 mm

𝑙𝑛 =length of leaf 𝑛

𝑙𝑛 + 1 = length of the leaf that is just shorter than 10 mm.

If the youngest leaf is exactly 10 mm young and it’s the fifth leaf, the PI will be 5. However, if
leaf 5 is just longer than 10 mm and a younger leaf is just shorter than 10 mm, the PI will be
somewhere between 5 and 6 depending on the length of 5 and 6.
Question: Calculate the PI for a plant with 20 leaves, the youngest of which is 7 mm and the next
being 15 mm.

Solution:

The number of leaves = 20∴ n = 19

Youngest = 7 mm

Next youngest =15 mm

log 15 − log 10
𝑃𝐼 = 19 +
log 15 − log 7

1.1760 − 1 0.1760
= 19 + = 19 + = 19 + 0.5317 = 19.53
1.1760 − 0.8450 0.3310

GROWTH HORMONES

It is now definitely known that certain chemical substances formed in very minute quantities as a
result of metabolism inside the plant body have profound influence on the growth of the plant
organs and on the various kinds of tropic movement exhibited by such organ. They also have a
significant effect on certain physiological processes. These substances are known as hormones or
growth hormones or phytohormones. Phytohormones include growth promoting and growth
retarding substances along with flowering, wound healing and vitamin substances.The major
classes of phytohormones are Auxin, Gibberellin, Cytokinins, Abscisic acid (ABA) and Ethylene.

Auxin: the term auxin was coined by F. W. Went in 1928. The term auxin literarily means arouser
and it was the first growth hormone discovered from plants. It is universally distributed in higher
plants as well as in lower plants (e.g. algae, fungi, bacteria, liverworts and ferns). The forms of
auxin that have been isolated include auxin a, auxin b and heteroauxin (indole acetic acid, IAA).
Auxins are widely distributed throughout the plant’s body but their distribution is more in actively
growing regions like shoot and root apices, lateral meristems, enlarging leaves, etc. Its
concentration is found to be more in shoot apex than root apex.Auxins are synthesized in the
meristematic regions of the plant and enlarging organs such as leaves, fruit aapices, seedling
coleoptiles and developing embryos and pollen. Auxin causes cell elongation and enlargement in
plants. Auxin also prolongs dormancy. Certain auxins are spread on potatoes to retard sprouting,
and thus auxin is very useful in the storage of fruits and vegetables. Auxin stimulates the formation
of adventitious roots and delay leaf abscission. Phototropism and geotropism occur due to unequal
distribution of auxin in the two faces of stem/root of the plant. In majority of vascular plants, apical
buds dominate over lateral buds. The growth of lateral buds occurs only after the removal of apical
bud. The phenomenon of suppression of growth of lateral buds by apical buds is called apical
dominance. Auxin enhances apical dominance.

Gibberellin: the hormone was first isolated by Yabuta and Sumuki in 1938 and named it
gibberellin after the name of the fungus from which they isolated it. Today, over 72 forms of
gibberellins are known out of which 15 have been isolated from Gibberellafujikuroi itself. Three
forms were extracted from beans, but the most common is the GA3 extracted from the fungus.
They are chemically known as gibberellic acid (GA). The hormone is widely distributed in nature.
In higher plants, they are found in the meristematic regions and young expanding organs, buds
embryos and seeds. The physiological effect of gibberellin is indirect. It exerts its effect by alerting
auxin status of the tissue. Gibberellins enhance longitudinal growth of internodes of genetically
dwarfed plants and breaks dormancy.

Cytokinins: Cytokinins was discovered by Miller et al. (1954). The first natural Cytokinins
discovered from plant iszeatin, isolated from maize seeds by Letham in 1963. It stimulates growth
by cell division. Cytokinins are widely distributed in plants either in free state or bound to
substances such as pentose sugar, inorganic phosphate or ribonucleotide. They are formed in the
root tips and transported upward via the xylem. Large amount of Cytokinins are found in
germinating seeds, bleeding sap, young developing fruits, endosperms and embryos.

Abscisic Acid (ABA): This hormone was isolated and identified simultaneously by a number of
scientists in different laboratories between 1863 and 1965. It was recognized as retarding hormone.
This substance is found in angiosperms, gymnosperms, pteridophytes and some mosses. Liverwort
and thallophytes do not have ABA. It is formed in the chloroplast of leaves but its formation is
also reported from other parts of plant. Fruits and seeds contain the highest amount of ABA. The
hormone accelerates senescence and abscission of leaves, flowers and fruits. Increased content of
ABA favours stomata closure. So, during water stress, its level increases in plant. It also induces
dormancy of seeds and buds. It acts to counteract the growth stimulating effect of auxin, gibberellin
and cytokinin.

Ethylene: this is the only hormone which is a gas at room temperature. It is produced in the fruit
and remains there. The existence of a gas which causes ripening of fruits was first observed in
1910 by H. H. Cousins. He observed that a gas emanating from oranges caused ripening of banana
stored together during shipping. But its nature was only established in 1934 by R. gane, and was
found to be ethylene, an unsaturated hydrocarbon. Ethylene is produced in fungi, bacteria and
higher plants. Ethylene mainly causes reduction in stem elongation and acceleration of senescence
in plants.

MOVEMENTS IN PLANTS

Living things are distinguished from non-living things by their power of movement. Protoplasm
is sensitive to various external agencies such as heat, light, electricity, gravity, chemicals, etc.,
which act as stimuli. Plants or plant organs often respond to such stimuli by moving in a particular
direction. The capacity of plants or their organs to receive and respond to stimuli is termed
irritability. Irritability is of benefit to plants by helping it adjust itself according to environmental
conditions. Water for the maintenance of turgidity, appropriate temperature, oxygen needed for
the respiratory release of energy, hormones and non-fatigue are essential factors necessary for
movement in plants.

Types of Movement

Plants display different kinds of movement which may be broadly classified into movement of
locomotion and movement of curvature. Movements of the protoplasm within a cell, free
movements of naked masses of protoplasm and those of unicellular or multicellular organs and
entire organisms are known as movements of locomotion, and these movements may be
spontaneous (autonomic) or induced (paratonic). Induced movements that occur as a result of an
influence by external stimuli are otherwise called taxism. Movements of locomotion are common
in lower plants.

Because higher plants are fixed to the ground, they are incapable of any locomotion. Some of their
organs however show different kinds of movement. The organs may move or change position or
direction by means of curvature. They move in order to get into position that will enable them
perform their functions more effectively. The movement of curvature can be broadly categorized
into variation movement and growth movement.

Variation Movements

These are movements of mature organs caused by variation in the turgidity of the cells of these
organs. The movements are fully reversible. Examples include sleep movements of leaves, touch
movement of the sensitive plant mimosa and the opening and closing of stomata.

Growth Movements

The movement of growth is the movement of growing organs and is due to unequal growth on
different sides of these organs. This kind of movement is slow and irreversible. Growth movements
are of three basic types:

– Tropic movement
– Nastic movement
– Nutational movement

Tropic Movement: tropic movement or tropism is unequal growth movement resulting from
environmental stimuli received directionally. The two most common examples are phototropism
and geotropism. Others are hydrotropism, chemotropism, thigmotropism, and thermotropism.

Phototropism or heliotropism is growth or movement towards or away from light. In positive

phototropic movement, growth is towards light while in negative phototropic movement, growth
is away from light. Runners and leaves that grow at right angle to the source of light are said to be
diaphototropic.

Geotropism is the growth movement in response to the force of gravity. Like phototropism, organs
of plant can be negatively, positively or diageotropic in their movement. The primary root of plant
is positively geotropic while the shoot is negatively geotropic. Lateral roots and branches are said
to be diageotropic.

Nastic Movement: this is the growth movement in which the direction of the response is
independent of the direction of the stimulus. For example, flower buds open in response to light.
Opening occurs because of a greater rate of growth in the upper surface than on the lower surface.
The result is curling of the flower bracts and the flower opens.
NutationalMovement: this is the growth movement resulting from unequal growth on the
opposite sides of the apex, and it is apparently independent of the environment. When a similar
pattern of growth covers the entire growing tip in a spiral manner around its long axis, the
movement is termed circumnutation.

JASPER
THE TALK ADDICT