UNIT SIX
PHYSIOLOGY OF PLANTS
Plants germinate from seeds, grow, develop, mature,
reproduce and die. Plant physiology is the study of
these processes. In other words, plant physiology
deals with how plants function. Plants are autotrophs
Chapter 1 and make their own food by utilising carbon dioxide,
water and sunlight. But before plants become
• PLANT-WATER RELATIONS photosynthetically competent after the emergence of
leaves, the germinating plants get their food from
Chapter 2 reserves in the cotyledons, as in peas and beans.
• PLANT NUTRITION As the plant grows, the role of food production is
shifted from cotyledons to fully exposed leaves, which
take a variety of shapes and sizes and carry out the
Chapter 3
process of photosynthesis. It is said that “only the
• PHOTOSYNTHESIS sun gives without taking.” And plants play a big role
in this process. The sun energy is converted to ATP,
Chapter 4 which is necessary for conversion of carbon dioxide
• RESPIRATION to carbohydrates. The carbohydrates (glucose, sucrose
or starch) are utilised in respiration by plants
themselves (and also by animals which eat plants) to
form ATP that provides the energy necessary for life’s
function. The process of respiration that produces ATP
uses oxygen that is released during photosynthesis.
It is this oxygen that changed the atmosphere from
one full of hydrogen to one rich in oxygen. Plants
possess the capability of taking up nitrogen and other
nutrients, along with water, from the soil through an
elaborate root system. The inorganic and organic
nutrients act as building blocks and help plants grow
and reproduce. All these activities in plants, like any
other living organism, follow the known chemical and
physical laws.
MELVIN CALVIN
(1911-)
Calvin born in Minnesota in April, 1911 received his B.S degree from
Michigan College of Mining and Technology (now Michigan Technological
University) and Ph.D. in Chemistry from the University of Minnesota.
He served as Professor of Chemistry at the University of California,
Berkeley.
He has served on many scientific boards for the U.S. Government.
Active in many professional societies, Calvin served the Royal Society
of London, which awarded him the Davy Medal in 1964 for his pioneering
work in chemistry and biology, particularly the photosynthesis studies.
His clear understanding of the nature of organic molecules proved
valuable in his subsequent work in biological chemistry. Investigations
of complex organic chemical systems using radio isotope C14 as a tracer
by his team were published as techniques in Isotopic Carbon (1949).
He along with J.A. Bassham studied reactions in green plants
forming sugar and other substances from raw materials like carbon
dioxide, water and minerals by labelling the carbon dioxide with C14.
These were described in The Path of Carbon in Photosynthesis (1957)
and The Photosynthesis of Carbon Compounds (1962). Calvin proposed
that plants change light energy to chemical energy by transferring an
electron in an organised array of pigment molecules and other
substances. This was substantiated by research not only in his
laboratory but also elsewhere later on. The mapping of the pathway of
carbon assimilation in photosynthesis earned him Nobel Prize in 1961.
The principles of photosynthesis as established by Calvin are, at
present, being used in studies on renewable resource for energy and
materials and basic studies in solar energy research. In addition, he
is also engaged in studies of chemical carcinogenesis studying the
molecular mechanisms of transformation of normal cells to malignant
cells and the synergism between chemical and viral carcinogenesis.
This study has indicated that chemicals act by triggering the integration
of oncogenic information already present in the cell.
Chapter 1
PLANT-WATER RELATIONS
Water is essential for all physiological activities the potential to do work. The potential
of plants. It acts as an excellent solvent and energy of water is referred to as water
helps in the uptake and distribution of mineral potential. That water has potential energy,
nutrients and other solutes required for growth can be exemplified by water stored behind
and development. Water also plays a direct role a dam. As this water runs downhill, its
in many useful reactions operating in cells. potential energy can be converted to
For example, water plays a key role in electrical energy by making the water fall
photosynthesis and acts as a source of oxygen. on turbines. Similarly, pressure is another
You know that cells are structural and source that provides energy to water. The
functional units of all organisms. In plants, increasing pressure increases the free
cells are composed of cell wall and protoplast. energy and, hence, the water potential in a
The term protoplast is used to refer collectively system.
to the plasma membrane and protoplasm. The
protoplasm refers to the living contents of cells Can we move the water up against
and consists of cytoplasm and a nucleus. It is gravity? Yes, we can, but only when
useful to consider the cell as having three pressure is applied by pressure. This
compartments : the vacuole, protoplasm and means that water moves from the point
the cell wall. The compartments are separated where water potential is greater, to the other
from each other by plasma membranes, the point where water potential is less. It can
tonoplast lying between the vacuole and the also be said that the difference in water
protoplasm, and the plasmalemma between the potential between two points is a measure
protoplasm and the cell wall. You also know of the amount of work (energy) needed to
that in many tissues, plasmodesmata connect move water from one point to the other.
the protoplasm of one cell to that of the next. Therefore, based on the concept of water
The plasma membrane is selectively permeable, potential, it is easier for the scientists to
which means it allows some materials to pass predict the way in which the water will
through it but not others. Also, a continuous move. Water potential is measured in terms
path for water exists from the soil through the of pressure. The common measurement
root, stem, leaf, and into the atmosphere. This unit of water potential is pascal, Pa (1
chapter covers absorption and movement of Megapascal, Mpa = 10 bars), and is
water within and between cells, its represented by the Greek letter, Psi (ψ).
translocation from root to leaf to atmosphere, It is measured in relative quantity, and
transpiration and stomatal movements. expressed as the difference between the
1.1 WATER POTENTIAL potential of a solution in a given state and
the potential of the same solution in a
All living organisms, including plants, standard state. Water potential is influenced
require free energy to grow and reproduce. by three factors : concentration, pressure,
In thermodynamics, free energy represents and gravity. Thus, water potential, ψw, of any
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4
solution can be represented as individual 1.2 ABSORPTION AND MOVEMENT
components as per the following equation : oF WATER
ψw = ψs + ψp + ψg Absorption of Water
Roots absorb water in plants. Root hairs present
Each component, ψs, ψp and ψg denotes,
on the root make a contact with soil particles
respectively, the effects of solutes, pressure,
and absorb water from the soil. Root hairs are
and gravity. Note that pure water has a higher
the outgrowths of the epidermal cells which
potential than the water inside a cell. increase the water absorbing surface area of the
Theoretically, ψw of the pure water is taken as roots. From root hair, the water moves through
zero and, therefore, ψw inside plant cells is the root cortex and to the xylem elements. Water
negative. in the root moves through two pathways :
The term, ψs, is called solute potential or (i) apoplast pathway, and
osmotic potential. Solutes present in a cell
(ii) symplast pathway.
(or in any solution) reduce the free energy of
water, or the water potential.
The term, ψp, is called pressure potential
or hydrostatic pressure of a solution. If the
pressure is positive, ψw will be raised and if the
pressure is negative, ψw will be reduced. The
positive hydrostatic pressure is called turgor
pressure and can keep on incresing till the cell
becomes fully turgid. At this stage the cell wall
prevents the protoplast from increasing in size
and exerts a pressure called wall pressure. The
ψp for pure water in an open beaker is zero.
The term, ψg, called gravity potential, denotes
the effect of gravity on water potential. It Fig. 1.1 The pathways of water movement
depends on the height of water above the
reference state of water, the density of water, The apoplastic movement of water occurs
and the acceleration due to gravity. If the exclusively through the cell wall without
vertical distances are small (less than 5 meters), crossing any membranes, while the symplastic
the ψg is negligible and, hence, ignored. movement occurs from cell to cell through the
In a plant cell, only ψs and ψp are important, plasmodesmata, as shown in Figure 1.1. The
and thus, the above equation is simplified as : symplast comprises the network of cytoplasm
of all cells inter-connected by plasmodesmata.
ψw = ψs + ψp Major proportion of water flow in the root
If the solute concentration is increased inside cortex occurs via the apoplast, as the cortical
the cell, the ψs is lowered, and thus, the ψw is cells are loosely packed, and thus, the cortex
offers no resistance. The apoplastic water
decreased. This allows water to move into the
movement beyond the cortex is blocked by the
cell from outside due to a water potential
casparian strip present in the endodermis
gradient. As water moves into the cell from
(Fig. 1.2). The casparian strip is composed of
outside, the hydrostatic pressure (ψp) increases. a wax-like substance called suberin, which
This results in an increased ψw of the cell, and blocks water and solute movement through
the difference between the inside and the the cell wall of the endodermis. Thus, beyond
outside ψw (∆ψw) is reduced. Water will move endodermis, water is forced to move through
out of the cell if it is placed in a solution that the cell membranes and cytoplasm (Symplastic
has lower ψs or ψw. pathway). This movement of water through
PLANT-WATER RELATIONS
5
thick
Fig. 1.2 Water movement from root hair to the cortex and to the xylem vessels, showing the apoplast
pathway and the cellular pathway through symplast and transmembrane. The casparian
strip present in the endodermis is shown in an enlarged view
the cell membrane is called transmembrane molecules is random and independent of
pathway. In this pathway, water may also cross each other.
through the tonoplast surrounding the Uptake and distribution of water, gases and
vacuole. Once the water reaches root xylem, solutes occur in the plants as a result of
transpiration drives the water to move to the diffusion, for example, supply of carbon dioxide
leaves through the stem. from atmosphere to the leaves for
Diffusion photosynthesis, and loss of water vapour from
Diffusion is a random movement of individual leaves to the atmosphere.
molecules from a region of higher concentration Permeability
to a region of lower concentration. Larger the The extent to which a membrane permits or
difference in concentration, more rapid is the flow restricts the movement of a substance is called
of molecules. Diffusion is more rapid in membrane permeability. It depends on the
gases than in liquids. When there is no net membrane composition, as well as the chemical
movement of molecules, a state of equilibrium nature of the solute. Permeability can be
is reached. In diffusion, the movement of measured readily by determining the rate at
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6
which solute passes through a membrane equilibrium is reached. Pressure can be
under a specific set of conditions. applied to the solution from the upper part of
Osmosis the funnel to prevent the movement of water
Diffusion of water through a semi-permeable into it through the membrane. The pressure
membrane is known as osmosis. The diffusion required to stop the movement of water
of water molecules continues to occur across completely is called osmotic pressure. It is
the membrane until an equilibrium is reached. also referred to as osmotic potential or solute
At equilibrium, the water potential on both potential. Osmotic pressure and osmotic
sides of the membrane is equal. Osmosis can potential are numerically equal, but osmotic
be demonstrated by a simple experiment, as potential has a negative sign. The magnitude
shown in Figure 1.3. of the osmotic pressure is a function of solute
concentration in a solution. It should be noted
that water not bounded by any membrane,
Pressure has no osmotic pressure. It has only the
potential to result into a pressure when placed
in an osmometer. That is why a solution has
an osmotic potential (ψs), which is the negative
of osmotic pressure (π; pi), that is :
h ψs = -π
If additional pressure is applied (more than
the osmotic pressure applied to prevent the flow
Sucrose of water into the solution in the funnel in the
solution
above experiment), then water can be made to
Membrane flow out of the solution into the water in the
beaker. This process is called reverse osmosis.
The process of reverse osmosis is used for
removing salts from saline water. Osmosis is
Water
driven by two factors :
(i) concentration of dissolved solutes in a
(a) (b)
solution, and
(ii) pressure difference.
Both these factors determine the chemical
Fig. 1.3 A demonstration of osmosis. A thistle potential of water, which is the driving force for
funnel is filled with sucrose solution water movement in plants. Water moves from a
and kept inverted in a beaker
region of high chemical potential of water to a
containing water. (a) Water will diffuse
across the membrane (as shown by region of low chemical potential of water. The
arrows) to raise the level of the solution chemical potential of water is also known as
in the funnel (b) Pressure can be water potential and represents the free energy
applied as shown to stop the water associated with water.
movement into the funnel
Plasmolysis
In this experiment, the water contained in If a turgid plant cell (cell with positive turgor
a beaker is separated from a solution in a pressure) is placed in a solution that has more
thistle funnel by a semi-permeable solutes, it exerts a higher osmotic pressure, and
membrane. (This kind of membrane can be water will move out of the cell by osmosis. This
obtained from an egg.) Water will continue to will cause vacuole and protoplast to shrink,
move across the membrane, resulting in the resulting the plasma membrane to fold inward
rise of the solution in the funnel until an and move away from the cell wall, as shown in
PLANT-WATER RELATIONS
7
(ψw = 0), a steep water potential gradient is
established. This allows rapid movement of
water to the surface of the dry seeds. The
imbibition of water into seeds continues until
an equilibrium between the water outside and
in the seed is reached.
1.3 THEORIES OF WATER TRANSLOCATION
How can water move to the top of a tree which
may be more than 100 m high? Different
theories have been put forth to explain this.
The three most prominent theories are :
(i) Root Pressure
(a) (b) (c)
(ii) Capillarity
(iii) Cohesion Theory
Fig. 1.4 Plasmolysis in an epidermal cell of a
Root Pressure
leaf. (a) cell under normal conditions Water from the soil is absorbed by root hairs
(b) cell when placed in a strong and is conducted through xylem vessels. If the
sucrose solution, plasma membrane stem of a plant is transversely cut above the
starts contracting (c) cell when placed soil surface, xylem sap will exude in the form
in a more concentrated solution loses of a drop from the cut surface. This indicates
lot of water and contracts much more
the presence of a positive pressure in the xylem.
This pressure is known as root pressure.
Mineral ions from the soil are taken up by
Figure 1.4. This is called plasmolysis. roots and get deposited in the xylem vessels.
Continuous loss of water from plant cells results Accumulation of ions leads to the lowering of
in wilting and drooping of leaves and stems. osmotic potential. This, in turn, results in
Imbibition lowered water potential in the xylem vessels.
Imbibition is a type of diffusion by which The low water potential of the xylem sap
movement of water takes place along a compared to that of the soil solution attracts
diffusion gradient. An adsorbant (for example, water, which is taken up inside the roots.
cellulose matrix) is required for the imbibition Remember, that water flows from a region of
to occur. Dry plant material or seeds act as high water potential to a region of low water
adsorbant to imbibe water and swell. The
potential.
swollen seeds produce a large pressure
developed by imbibition. This pressure makes Capillarity
the seedlings to emerge above the ground The uptake of water through xylem vessels is
through the soil surface. The pressure, called also possible in small-sized plants through
imbibition pressure, is also known as matric capillarity. Capillarity refers to a rise in water
potential (i.e. water potential of the matrix) in in tubes of small diameter kept in a vessel
the context of plant-water relations. containing water. This rise in water is due to
Two conditions are necessary for the
the forces of adhesion and cohesion. Adhesive
imbibition to take place. They are :
forces attract molecules of different kinds,
(i) water potential gradient between the
surface of the adsorbant and the liquid whereas cohesive forces attract molecules of the
imbibed, and same kind to each other. Force of gravity also
(ii) affinity between the adsorbant and the affects water uptake by capillarity.
imbibed liquid. According to this theory, water is first taken
Dry seeds have a highly negative water in due to the force of adhesion between water
potential. When these seeds are placed in water and the walls of thin xylem vessels (tracheids).
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8
As the water flows upward along the wall, strong forms a continuous water column from roots to
cohesive forces between water molecules come the leaves due to cohesive and adhesive
into play to pull the water upward. This upward properties of water molecules. Also, the walls of
pull of water continues until the forces of xylem vessels, made up of ligno-cellulose, have
adhesion and cohesion are balanced by the strong affinity for water molecules. The water
downward force of gravity. potential of root cells being lower than the water
Cohesion Theory potential of soil, movement of water occurs from
Cohesion theory is the most accepted theory of soil to the roots. Uptake of water, as described
water movement through plants. This theory by the Cohesion theory, is also referred to as
was given by Henry Dixon in 1914. As the name transpirational pull (Fig. 1.5).
suggests, the Cohesion theory is based on the 1.4 TRANSPIRATION
force of cohesion between water molecules. This
allows formation of a continuous water column Loss of water in the form of water vapour from
throughout the plant, that is, from the top of the the plant to the atmosphere is known as
plant down to the root tip. According to this transpiration. About 90 per cent of the total
theory, evaporation of water from the leaf to the transpiration occurs from leaves. Leaf surfaces,
atmosphere results in a decrease in the water both upper and lower, contain small pores, called
potential of the epidermal cells, which are in stomata, through which water is transpired to
direct contact with the atmosphere. The water the atmosphere. The stomatal pores open into
lost from these cells is replaced by the water the intercellular spaces of the leaf and provide
moving from the adjacent cells along a water an uninterrupted path from the interior of the
potential gradient. The water lost from leaf cells leaf to the external environment (Fig. 1.6).
is, responsible for generating a negative ψw in Significance of Transpiration
the xylem elements creating a tension or If the transpirational loss of water is higher
transpiration pull. As a result water movement than the amount of water absorbed by the roots
from the soil, plants undergo water stress. This
results in decline of growth in plants. The
question is, then why transpiration is
Transpiration important and what is its significance.
Evaporation
Transpiration helps in the movement of xylem
sap, which simultaneously increases the
absorption of mineral nutrients by the roots
Porous from the soil. As the evaporation causes cooling,
pot
it assists in the cooling of leaves. Not all the
solar radiations absorbed by leaves are used
Water
in photosynthesis, but some will cause
warming of the leaves. Transpiration, however,
reduces the heating of the leaves.
Mercury Factors affecting Transpiration
T ranspiration is af fected by three
environmental factors, that is, humidity,
temperature and wind speed. These factors
influence the rate of water diffusion from the
leaves to the atmosphere.
(a) (b) If the humidity is high, water is lost slowly
into the air as a result of transpiration and vice
Fig. 1.5 Demonstration of Cohesion theory of versa. The rate of evaporation doubles with
water translocation (a) Evaporation; every rise in temperature by 10°C. As
and (b) Transpiration transpiration causes cooling of leaves, the leaf
PLANT-WATER RELATIONS
9
Fig. 1.6 Water movement through the leaf to the atmosphere in the form of vapour
temperature does not rise as much as that of 1.5 OPENING AND CLOSING OF STOMATA
the air. High temperature, influencing the All leaves have a large number of tiny pores,
closure of stomata, reduces transpiration. called stomata. These stomata are located on
Wind speed also has an influence on the rate the surface of the epidermal layer of leaves.
of transpiration. A high wind speed or a dry Each stoma is composed of two special types
breeze will greatly increase transpiration. of epidermal cells, called guard cells. The
Some other plant factors also affect the rate guard cells control the opening and closing of
of transpiration. These factors include : efficiency stomata, which, in turn, regulate the loss of
of water uptake by roots, the leaf area and the water from the leaves to the atmosphere
leaf structure. Larger leaf area will transpire through transpiration. The size of a typical
more water into the atmosphere. Leaves of plants stomatal pore ranges from 3-12 µm in width
which grow in a dry habitat develop structures and from 10-14 µm in length. The number of
like thick cuticle and sunken stomata (deep- such tiny pores varies from 1,000 to 60,000
seated) to cut down the loss of water through per cm2 of the leaf surface. Stomata are found
transpiration. on the lower (abaxial) as well as upper (adaxial)
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Table 1.1 : Number of Stomata on monocot leaves (grasses), the stomatal density
the Upper and Lower or number is almost the same on both the
Surfaces of Leaves surfaces. Leaves of dicot plants usually contain
Plant Number of Stomata/mm2 fewer stomata on the upper surface.
Upper Surface Lower Surface Opening and closing of stomata is controlled
by accumulation of solutes in the guard cells.
Monocot Solutes are taken in by the guard cells from the
Wheat 50 40 neighbouring epidermal and mesophyll cells.
Barley 70 85
As a result, both osmotic potential and water
potential of the guard cells are lowered. This
Onion 175 175
would create a water potential gradient between
Dicot the guard cells and the neighbouring cells, and
will make the water move into the guard cells.
Sunflower 120 175
The guard cells become turgid, and swell in size,
Alfalfa 169 188 resulting in stomatal opening. With a decline in
Geranium 29 179 guard cell solutes, water moves out of the guard
cells, making them flaccid. As a result, the
stomata close. Thus, the opening and closing
surfaces of the leaf. However, the number of of stomata is controlled by osmotic movement
stomata on the upper and the lower surfaces
of water in or out of the guard cells along a water
of leaves varies among species (Table 1.1). In
potential gradient.
The major solute, which is taken in by the
guard cells from the neighbouring cells, is
potassium. The rise in potassium levels
causes stomatal opening and its decrease
Vacuole stimulates stomatal closing. The uptake of
H+ H+
potassium controls the gradient in the water
ATP potential. This, in turn, triggers osmotic flow
K+ K+
of water into the guard cells raising the turgor
pressure. Role of K+ in stomatal opening is
Cl– Cl–
now universally accepted. The extent of K+
2H + accumulation in guard cells determines the
CO 2 Malate size of the stomatal opening.
The accumulation of large amounts of
PEPcase
potassium (K+) in guard cells is electrically
balanced by the uptake of negatively charged
PEP Starch ions – chloride and malate (Fig. 1.7). The high
amount of malate in guard cells of open
Guard cell stomata accumulates by hydrolysis of starch.
Factors affecting Stomatal Movement
Many environmental factors affect stomatal
Fig. 1.7 Role of potassium, chloride and movement. Most prominent factors include
malate ions in stomatal opening. The light, temperature, water availability to plants
ions accumulate in the vacuole of
and CO2 concentration. Some endogenous
guard cells, lowering the water
potential and thereby increasing factors, like K+, Cl– and H– ions and organic
water uptake and subsequently acids, also influence stomatal movement.
opening the stomata. (PEPcase = Light : Stomata open in the presence of
Phosphoenol pyruvate carboxylase) light and close in darkness. Light intensity
PLANT-WATER RELATIONS
11
required to open the stomata is very low, as called water deficit or moisture deficit)
compared to the intensity required for induces stomatal closure. This happens to
photosynthesis. Even moonlight is sufficient to conserve moisture by plants by cutting down
keep the stomata open in some plant species. the transpirational loss of water.
In plants with Crassulacean Acid Metabolism CO2 concentration : With an increase in
(CAM), stomata open during dark and remain carbon dioxide concentration inside the leaf,
closed during the day (see Chapter 3). This the stomata close. This happens even under
unique behaviour of stomata is a kind of the light. In some plant species, stomata also
adaptation to conserve moisture in CAM plants, close if we merely breath on their leaves. It is
for example, pineapple, agave, etc. the internal leaf carbon dioxide concentration
Temperature : Stomata tend to open more
rather than the atmospheric carbon dioxide
with an increase in temperature and close with
that dictates stomatal opening. If plants
a decrease in temperature. In some plant
are transferred to carbon dioxide-free
species, stomata remain closed even under
environment, but kept in darkness, the
continuous light at 0°C. However, if the
stomata will still remain closed. This means
temperature is increased, stomatal opening in
these species increases. At temperatures higher that, since the internal carbon dioxide is not
than 30°C, there is a decline in stomatal opening utilised due to absence of photosynthesis in
in some species. the dark, it influences the stomata to remain
Water availability : If the water available closed. However, if these plants are exposed to
to plants is less and transpiration rate is high, light, photosynthesis will utilise internal CO2,
plants undergo water stress. Water stress (also permitting the stomata to open.
SUMMARY
Water is extremely essential for plant growth and development. It is absorbed
by the root hairs from soil and translocated to the leaves. Various mechanisms
operate for the water to move from one cell to another and, therefore, from
root to stem to leaf to the atmosphere. The movement of water in the roots is
through apoplast pathway and symplast pathway. However, major portion of
water in the root flows via the apoplast.
Different theories have been put forth for explaining water translocation
within a plant. The three most important theories are root pressure, capillarity
and Cohesion Theory. Most of the water taken up by plants is lost to the
atmosphere by transpiration, which occurs through tiny pores called stomata,
located on the leaf surface. Only a fraction of water absorbed is utilised for
the purpose of growth, maintenance and reproduction of the plants. A large
number of stomata are present on the leaves, mostly on the lower leaf surface.
The stomatal movements, i.e., opening and closing, control the flow of water
from the plants to the atmosphere through transpiration. Transpiration helps
in the movement of xylem sap, and simultaneously increases absorption of
mineral nutrients by the roots from the soil. It also assists in cooling by
evaporation of water and thereby reducing heat load on the leaves. Various
environmental factors contribute to the occurrence of transpiration and
control of stomatal movements. These factors mostly include light,
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temperature, humidity, CO2 concentration, and water availability. In addition,
many endogenous factors also influence both transpiration and stomatal
movements.
EXERCISES
1. Describe osmosis as a special case of diffusion.
2. Distinguish between the following :
(a) Osmotic pressure and osmotic potential
(b) Osmotic potential and matric potential
(c) Diffusion and osmosis
(d) Osmosis and imbibition
3. Describe the theories related to translocation of water.
4. In what way does the concept of water potential help in explaining
water movement?
5. Of what importance is potassium in the opening and closing of
stomata?
6. Is there a general mechanism to explain the opening and closing of
stomata? Justify your answer.
7. Mention some of the factors that influence stomatal opening and
closing. How are these factors involved in regulating stomatal
behaviour?
8. Does transpiration serve any useful function in the plant?
9. Write short notes on
(a) Plasmolysis
(c) Imbibition
(b) Osmosis
(d) Root pressure
10. Describe the role of osmotic potential in regulating water potential of
plant cells.