UNIT 11 PLANT WATER RELATIONS

Structure
Introduction
Objectives
Plant Water
Early History of Ascent of Sap
The Pathway of Transport of Water
Some Basic Physical Concepts
Resistances to Water Movement and Water Flux
Gradients of Water Potential
Water Absorption
Water Loss
Stomata
The Mechanism of Stomatal Open~ng
Factors Controlling Stomatal Aperture
Summary
Terminal Questions
Answers

11.1 INTRODUCTION
At one time it was thought that man would be able to colonise the moon and other
planets. However, even before man landed on moon his hope was shattered. The
reason, neither there was the atmosphere containing gases which sustain us on the
earth nor was there any water which is essential for life. Life originated in water of
the oceans and water remains a key molecule in maintaining life on earth.
The intimate relationship between water and plant is apparent from the greenery in
areas where water is abundant and barren deserts in areas where it is in extreme
deficit. Among the environmental factors - light, temperature, water and soil, it is
the water that limits plant growth in virtually all environments.
The quantity of water absorbed by plant is enormous and is far greater than taken
by animals of the same weight. This is because the loss of water by plants through
transpiration is enormous. A major challenge before plant physiologists is to find
ways to decrease water losses and increase the efficiency of water use by the plants.
Plants can move water to great heights in trees. Even the tallest tree Sequoia
sempewirem ([Link], and still growing) found in California faces no problem in
moving water to its top leaves. This was puzzling for the scientists. They tried to find
out the forces that move water to such great heights.
We have the following questions before us with regard to water relations of plants)
i) What is the force that drives the flow and direction of water in plants? ii) What
are the forces that drive water to great heights in trees? iii) What are the factors that
control the opening and closure of stomata1 aperture? and iv) Why do plants
transpire so much water in an apparently wasteful way?
In this unit we will try to find answer to these questions.

Objectives
deqcribe early experiments on water movement through vascular plants,
explain the cohesion-tension theory of water movement in plants,
draw and explain radial movement of water from soil to roots and long distance
transport from xylem to leaves,
describe the factors that affect water potential and explain their significance in the
trans$rt of water in soil-plant-atmosphere system.
explain how differences in water potential, A$,, affect the direction of water
movement,
calculate I),, Sip, A*,, water flux and resistance using mathematical expression;
Plant Physiology-I discuss varlous resistances that impede water flow in plant and explain their
significance to the plant, ,
describe the factors that affect water absorption and water loss,
relate the structure and properties of stomata to-their function,
explain how changes in turgor bring about opening and closure of stomata,
explain the causes that alter the relative turgor in the guard cells, and
list factors that control movement of stomata1 aperture.

11.2 PLANT WATER

Role of Water in Plants
You know that water is the main constituent of plant cells. It performs the following
major functions.
i) Water as a Solvent
Water is a very good solvent. It easily dissolves electrolytes and [Link] such
as glucose and amino acids. As you know life evolved as a result of chemical reactions
that occurred in aqueous medium of the oceans and even today we know that all
reactions in a cell occur in aqueous medium.
ii) Water as a Chemical Reactant
Water participates in many biochemical reactions. It is involved in photosynthesis,
the most important process of life. The photochemical splitting of water ebolves
oxygen, and the hydrogen atoms are denotated to C 0 2 for making glucose. D u ~ g
catabolism, carbohydrates, fats and proteins require water for hydrolyses. In the
cpurse of your study, you will come across many other biochemical reactions which
also involve water.
iii) Water Provides Turgidity to the Cells
Plants maintain their shape due to turgidity which is brought about by the hydrostatic
pressure of water in the cells. If water moves out, the cells become flaccid (Fig. 11.1).
Hydrostatic pressure is also necessary fbr the'enlargement of cells and as a
consequence growth results.

~ .:CeUs of a normal (a) and a wilted plant (b). In fully turgid plant the central vacuole of&
l ~ i 11.1 cell
is filled with water and the protoplam is pusbed towards the d wall, sbwclhg it tight, whik
in the wilted plant the cell walls may partially deflate atter the vacuole shrinks.

How much Water is Present in Plants?

The amount of water in plants varies among different species depending upon their
structure. Aquatic plants such as Chlamydohtonas, Spirogyra, Chara, red algae and
others contain 97-98% of their weight as water. This is true also of Azolla, Eichhomia
and other fresh water plants. However, among terrestrial plants, whether they are
crops such as wheat, rice, maize, or trees such as sheesham, mango, neem and others,
the amount of water varies in different plant parts. It would be as high as 95% or
more in young roots and as low as 30-40% in wood of a tree trunk. Young leaves
often contain 85-90% water whereas mature leaves contain 60-80%. This means that
the amount of water changes during the growth of the plant as well as d u r i n i e
growth of an organ such as leaf and seed.
Relation of Water Content to Functions Plant Water Relations

If an aquatic plant appearing turgid is removed from water and left in the open, it
quickly wilts. The water content of the leaves may change hardly by 3 to 5 per cent
but the plant looks functionless. This is also true for young leaves of wheat which
have 90-95% water, they wilt if water content drops by 3-4 per cent. However, an
older leaf has only 80-85% water but is fully turgid. Thus, water content cannot be
the basis of judging the activity of leaves. Therefore, it is necessary to have an
expression for plant water status which could be related to plant function. We will Water has a peat tensile strength
discuss this in one of the sections later. Theoretical calculations, from
heats of evaporation and surface
tension, indicate a tensile strength
for water of several thousand
11.3 EARLY HISTORY OF ASCENT OF SAP atmospheres. Experimental
values, however, are somewhat
It is a familiar fact that water runs downhill. But plants can raise water upwards from lower, ranging from 25 to 300
soil by roots to great heights of trees. This was one of the puzzles of early plant atmospheres. In one illustrative
experiment the British
physiologists. They tried to know what the forces are that drive water upwards. They investigator H.M. Budgett wrung
considered two possibilities : i) water is either pushed up by the driving forces that two polished steel plates together
might develop at the bottom (roots) or ii) it is pulled up by the forces created at the with a film of water between
top (leaf) of the plant. them. Tensions of up to 60
kilograms per square centimeter
Even 400 years ago, it was recognised that it is the xylem conduits and not the phloem was required to pull the plates
that translocate water in plants. It was Italian anatomist, Marcello Malpighi who in apart.
1679 demonstrated that if a ring of phloem was removed from the stem the path of
water was unaffected. Water moves through dead xylem cells was shown by Edward
Strasburger in 1883 who sawed an oak tree and placed it in a bucket of water
containing picric acid or CuS04. Though the chemical killed the bark and other living
cells the movement of water was uninterrupted. Experiments have also been done
ing coloured or radioactive water. The water is observed to move into the root
and upward through xylem. Now, we know that xylem vessels form an
plumbing network whose supply lines extend to all parts of the plant.

(a) Simple suction (b) Root pressure (c) Capillary action

I FLg. 11.2 :Experimentson water movement through vascular plants. a) Simple suction -mercury is raised
76 cm up In an evacuated tube by atmospherk pressure. b) Root pressure-a glass tube Is fitted
on the cut stem of a potted plant. The soU is kept weil watered. Root pressure forces water to Mercury
exude out from the cut stem and up the tube. The water column may rise up to a height of
14 m or more. c) Capillsry - water rises In the capillary tube due to high surface tenslon Fig. 11.3 : Demonstration of the
of water. The greeter tbe surface tension greater the rise in the capiUary. principle of water movement up
the plant stem by a physical model.
For investigating which of the forces were responsible for the upward movement of Evaporation of water from the
water, initially forces such as atmospheric pressure, capillary action and root pressure porous clay pot exerts more pulling
were considered (Fig. 11.2 a, b and c). But none of these forces could account for 'force than simple suction. It can
the movement of water beyond a height of 100 cm. It was an Irish Botanist Josef ' pull mercury behind it to a height.
Bohm (1883) who demonstrated by a simple experiment that if water were to be of more than 100 cm.
evaporated from a dosed system like a porous pot connected to a set up as shown in
Fig. 11.3, mercury could be lifted to a height of more than 100 cm, considerably
 higher than 76 cm to which it can be pulled by a vacuum. Irish Botanist H.H. Dixon
and his collaborator J: Joly repeated the same experiment using a transpiring pine
Guttation
Under certain environmental leaf instead of a porous pot and got the same results. A similar experiment shown in
conditions root pressure forces Fig. 11.4 illustrates the rise of water in plants. Dixon and Joly (1895) then proposed -.
water through special water pores [Link] famous cohesion theory of 'mcent of sap.
around the edge of the leaves or
at the tip. This is known as .I I

guttation.
Leafy shoot
I

Fig. 11.4 :Experiment to show the cohesion-tendon theory of water movement in the p h t . Two long ulendcr
tubes dipped in water from below are extended up to the height of tk W i n g . A v a x ~ u mpomp
fails to suction the water to this height while a tiny leafy shoot succeeds. A short piece drubber
tubing is used to connect the cut tip of a leafy shoot to a slender glass tubing (14m length m d
0.5 m m in diameter) which is dipped in a beaker containing water and dye. The entire system h
made continuous water filled air proof system.

You are aware that plants lose water through stomata1 openings by the process of
transpiration. Dixon's theory suggests that water can be continuously pulled upward
due to the transpiration pull. As the water molecules vacate transpiration sites (the
mesophyll cells of leaf lining stomata) due to evaporation, other water molecules are
pulled to fill their places. This results in the creation of a negative hydrostatic pressure
downward extending from leaves to the xylem vessels and finally to the roots. In otlier
words, a state of water tension is created inside the xylem conduits. The water
molecule$ do not snap away from each other due to tension in the xylem, instead
Fig. 11.5 : Diagrammatic they travel as a continuous column due to their cohesive property. Cohesion is a
representation of the cohesiveness phenomenon by which water molecules cling tightly to one another by hydrogen
of water molecules due to hydrogen bonds (Fig. 11.5) and resist being pulled apart. If water is indeed moving through an
bonds. Water molecules cling upward pull, it would tie expected to recede in the xylem, if its continuum is broken
together due to electrostatic by cutting a branch above the ground level. The state of water in xylem is more or
attraction between the partial
negative charge on the oxygen less analogous to a stretched rubber band. Suppose if we cut a stretched rubber band
atom of one water molecule and at some point, it would immediately recede in both direction from the point where
the partial positive charge on the it is cut. [Link] receding occurs in xylem vessels and water pulls away from the
hydrogen atom of an adjacent cut point. The pulling force of water column or tension in the xylem can be measured
'water molecule. by a Scholander pressure bomb (Fig. 11.6) by forcing the xylem sap back to the
original point by applying pressure.
 Plant Waler Relalions

Xylem sap forced back, .

to original pmnt

Pressure gauge
gai delivery

Water in xylem pulled

back to here

Fig. 11.6 :Demonstrationof tenslon in xylem sap. When a twig is cut the tension in xylem ptllls the sap away
from the cut point, The pressure applied in Soholander Pressure Bomb forces xylem sap back to
the cut point.
It is important to point out here that the dead cells of the xylem are greatly
strengthened with cellulose fibrils encrusted with lignin. Such strengthening of walls
can be compared with reinforced concrete and thus the xylem cells are strong enough
not to collapse when water is pulled through them.
Studies on the movement of water at various times of the day also suggest that the
"motor" for ascent of sap lies in the leaves of the tree. The velocity of sap flow in
the wood was also measured by inserting a heating element into the xylem. After a
time interval the distance covered by the heated xylem sap was measured by a
thermocouple. The velocities as high as 75-100 c m h r were recorded (Fig. 11.7).
Although, the cohesion theory of wdter was proposed about a century ago yet it holds
1 good even today. However, it explains only the long distance transport of water

I through dead xylem tissue. Many qdstions -such as what the factors are that affect
water absorption from soil, how does water move from cell to cell and in tissues other
than xylem and what are the driving forces that determine the direction of water
movement - remain to be answered.
Fig. 11.7 :Measurement of velocity
of xylem sap. A small heating
element can be ihserted at a point
In the following sections we explain to you the concept of water potential which can in the t ~ n upto
k the xylem. After
1
tell us not only the water status, but also the direction and rate of movement of water a time interval the upward
*
in a plant. movement of the sap can be
detected by a thermocouple. ,

11.4 THE PATHWAY OF TRANSPORT OF WATER

Plants absorb water from the soil by roots, mainly near the root-hair zone in the
region of maturation (Fig. 11.8). The radial movement of water is shown in

. (a)

pa. 11.B :a) General morphology of root tip ahowing root cap, merBtematic zone, zone of elongation and
zone of maturation with root hair. b) position of various tissues including xylem and phloem.
Plant ~ y s i o l o w - 1 Fig. 11.9a. At most stages of its journey water molecules have the option of moving either
through protoplast or through cell walls. Water may first enter the epidermal cells of
the root by crossing plasma membrane and then move along through the cytoplasm
of the cortical cells, the endodermis, the pericycle and finally into the xylem vessels
andlor tracheids. The movement of water through cell to cell occurs probably via
plasmodesmata - the protoplasmic bridges between the cells. This transport route
is termed symplastic pathway (Fig. 11.9b). Since cells are joined via intercellular
connections, the entire living portion of the plant forms a continuous single entity
and is called symplasm.

The other route for the transport of water is through cell walls, intercellular spaces
and non-living cells of the xylem (Fig. 11.9b). This is called apoplastic pathway and
the non-living portion of plants is called apoplasm. The cell wall is composed of
hydrophilic substances like cellulose, hemicellulose, pectin, lignin and hydrophilic
proteins, and carbohydrates, polymer<of gums and mucilages. The molecules of
these substances retain a lot of water and let it permeate easily without any resistance,
when the supply of water is in plenty. When water moves through the cell walls of
epidermis and the'cortex, its moveyent is restricted at the highly packed endodermal
cells, because the radial and transverse cell walls but not the tangential walls of these
cells are lined with suberin a hydrophobic substance in the form of strip called
Casparian strip (Fig. 11.9~).
'
. Chemically suberin is like cutin and lignin of cuticle which are impermeable to water.
Therefore, water crosses membranes in order to enter endodermal cells and joins the
symplastic route. Some water may enter cells at any point prior to endodermis and
&

Casparian
Cortex strip
I I

Casparian strip
\ \

Outside of Casparian strip

Transverse wall
a1 wall

Fig. 11.9 : a) Cross-section of a root showing radial movement of water from root to xylem. b) symplastic
and apoplastic routes. c) Casparian ships.
 Plant Water Relations
I join the symplastic pathway. Once the water reaches xylem conduits it spontaneously
moves upwards through xylem of root, shoot, petiole and finally in the tracheid of
/ leaf vein and mesophyll cells (Fig. 11.10). Most of the water evaporates from the
mesophyll cells that line the stomatal cavity and diffuses out to the air through
i stomatal pore.

3.0 2
2
. -
s
.-bCd .
I-'

Y
0
a

Fig. 11.10 :Tbe pathway of water movement in a plant : a) whole plant. b) movement through xylem. The
-
scale shows water potential at different points in the plant.

SAQ 1
a) Which of the following statements are true.:

i) A plant (without roots) would not translocate a dye solution

if first placed in picric acid solution for about one hour.

ii) If a rapidly transpiring plant Is cut off just above the ground,
water will ooze out of xylem vessels.
I
/
iii) Removing all the leaves from a plant will probably reduce
flow of water up the stem.

iv) A soap solution injected-in the xylem of a tree-trunk could

prevent water from reaching at the top.

Hint : Soap solution reduces the surface tension of water.

 11.5 SOME BASIC PHYSICAL CONCEPTS
The main idea of this section is to explain the concept of chemical potential of water
or water potential and the effect of various factors that change its value in plant cell
and its immediate environment. The flow of water depends upon the gradient of
water potential in soil-plant-atmosphericsystem. The understanding of this section
requires some knowledge of certain basic physical concepts such as diffusion,
osmosis, imbibition, and chemical potential. We will explain them briefly. (You need
to revise Section 7.3 of Cell Biology Course before reading this section.)

Diffusion
Diffusion is a spontaneous process that leads to net movement of a substance from
a region of higher concentration to a region of lower concentration. 1"tcan also be
defined as the net movement of molecules from region of high free energy to a region
' of low free energy.
The principle of diffusion is of importance in plants because several of the transport
processes such as uptake of water and minerals, intake of C 0 2and release of O2by
leaf cells, loss of water due to transpiration depend in part on diffusion.
Osmosis
Osmosis is a special case of diffusion. Here, the movement of water (solvent) takes
place from a region of higher concentration to a region of lower concentration, if the
two are separated by a semi-permeable membrane.
Osmotic Pressure t

It is the pressure necessary to prevent the fiow of water or a pure solvent to a solution
(osmosis) when the two are separated by a semi-permeable membrane. Osmotic
pressure is given by the following equation
n = CRT ... (11.1)
IT =. osmotic pressure in bars

C = concentration of solution in moVl

It is the pressure eierted by R = gas constant litre bar m o l - ! ~ ~ '
vapdur over a liquid where it is in
equilibrium with itself. Its value is 0.08 litre bar mol-'K-'
T = absolute temperature in Kelvin
The osmotic pressure of a molar solution (i.e. one moleflitre) is thus
-
- - X1 mol 0.08 litre bar 273
litre mol K
= 21.84 bars (T = 273 K)
Imbibition
It is the process of absorption of water to the nearly dry surface such as seeds and'
wood. There is liberation of substantial amount of heat during the process.
Gradient
It is the difference between concentration or pressure or any other parameter
indicative of energy between two specified points. The direction of flow is always
from higher energy towards lower energy.

Chemical Potential
The chemical potential of a substafidk a system is a measure of its free energy, i.e.
.energy available to do work. Here the system refers to thermodynamic concept
wherein studies are of system rather than individuals or bodies. Thus when we study
the-propertiesof a solution in the container, we are studying a system wherein each
individual component interacts internally. For example, suppose we have pure water
in a system be it in a beaker or soI1,'all the molecules of water can do work, so their
free energy is maximum. When a small amount of sugar is added a few molecules of
water get associated with each molecule of sugar,.the free energy of water decreases.
Hence, the free energy of pure water is always maximum. Addition of any solute
lowers the free energy or chemical potential of water.
 Plant Water Relations
The chemical potential of water in a solution is rdated to its vapour pressure and is
given by the following equation.

h - p w O = RTIn
eO
AF = RTIn ... (11.2)
eO .
F, = chemical potential of water in question (joules/mole)
pwO= chemical potential of-pure water under STP
A F = change in free energy
R = gas constant
T = absolute temperature
e = vapour pressure of water in question
eP = vapour pressure of pure water
Note that
Relative huniidity = x 100
eO

If e is also pure water then In is zero and A F becomes zero. So the chemical
eO
potential of pure water is set to zero. If e is less than that of pure water then In 2
eO will
be negative number, therefore, AILwill be less than zero. - a negative number.

Water Potential .
It is the difference between chemical potential of water at any point in a system and
that of pure water at STP. By convention the chemical potential of water is referred
to as water potential and is denoted by +,(+-pronounced as psi).
+, is expressed in pressure units. It is the free energy per unit volume of water (joules
\
per cm3).

. I

G = partial molal volume

If water potential of the source (the region supplying the water) is higher than the partl~lmom (J) of a
water potential of the sink (the receiving region), then there is a spontaneous transfer solution is the change in volume
of water from source to sink (+, ,,,, > +,,,J. of a solution when one mole of a
substance is added to it.
By convention the water potential of pure water is taken as zero. Therefore, water
potentials other than that of pure water will generally be negative. Thus lower
potential means a more negative value, and higher potential a less negative value.
Water potential in plants is affected by the solute, hydrostatic pressure and matric
forces. In order to predict the movement of water inside or outside a cell we must
consider the effect of the above three factors.

Effect of Solute
Let us take pure water in two chambers A and B separated by a semi-permeable Water potential is expressed in
membrane (Fig. 11.11a). The water potential of both the chambers is zero. Addition pressure units. Energy per unit
of solute in chamber A (Fig. 1 l . l l b ) would reduce the free energy of water and the volume of water is expressed in
water potential will fall below zero. Consequently, water will move from B to A (Fig. joules per cm'. [Link] is equivalent
to dynes per cm2.lo6 dynes = 1
11.11~).The effect of dissolved solutes on water potential (JI,) is called osmotic bar. The present unit used for
potential (JI,). It can be estimated numerically if we know the osmotic pressure of pressure is pascal (Pa). It is a
the solution. The two are related as pressure equal to the force of one
,. Newton acting dniformly over one
IT=-IT
... (11.3) square meter.
For example, if n of a solution is 5 bars then JI, would be -5 bar.
,1 bar = 16Pa
Effect of Pressure ld Pa = 1 KPa (1 kilo pascal)
* lo3KPa = 1 MPa (mega Pascal)
Let us now see the effect of pressure on water potential. As Fig. l l . l l d illustrates,
when pressure is applied the flow of waterFbegins from chamber A to chamber B 13
through a semi-permeable membrane. This means that pressure increases the free
energy of water and thus raises the potential of pure water above zero. The effect of
pressure on water potential is called pressure potential and is designated as JI,. The
level of water in B rises due to increase in water potential of A (Fig. [Link]).
What would happen if pressure is applied on the chamber containing solutes? Now
the +, will be affected by both solute and pressure. The solute would lower the water
potential and pressure will raise the water potential. So the flow of water from B to
A would start decreasing. An equilibrium condition will reach when +,will be equal
but opposite in magnitude to +,
and there will no net flow of water in the two
chambers (Fig. 11.1le). This can be represented by the following equation:
+w(B) = ($T + JI,)B ... (11.4)
Let us suppose if JI, is equal and opposite to JI,. Then
JIWA = ($,A + JImB)

d e f
Fig.11.11 :Experiment to show the effect of solute and pressure on water potential, see text for details.

Effect of Matric Pressure

Water can get absorbed to the wettable surface of solids such as soil, wood, seeds
and cellular constituents. A force operates between solid-liquid interface and is called
matric suction or matric pressure. The absorption process is accompanied by heat loss
and results in decrease in free energy of water. In other words, the effect of matric
forces on water potential is called matric potential (JI,). Its value will be negative.
In a well watered soil JIiis not very significant, however, when the soil is near drying
+,, determines water potential of the soil. .

So we find that the JI, is composed of the following main component forces:
JIw=JI, + JIp + JIm + JI... ... (11.5)
JI.,. = any other forces that may inflhence JI,.
 Plant Water ~elatlom
11.6 RESISTANCES TO WATER MOVEMENT AND
WATER FLUX
We have explained earlier that if water potential drops from source to sink (I),,,, >
than there will be spontaneous flow of water. But we do not know the rate of
this transfer i.e. flux.
. Recall from Section 7.2 of the Unit 7, Cell Biology Course, the rate of diffusion dcldt
is flux. The flux for water flow is denoted by Jwwhich is volume of water flow through
unit surface area per unit time. Water in plants flows from cell to cell and also through
cell walls. -
When water moves from cell to cell the flow is the function of water permeability of
the membrane. The flux of water is given by
Jw =LpAI), ... (11.6)
L, = permeability coefficient of limiting membranes
AI), =. difference in water potential at two points

From equation (11.4) we know that

= (I), + I)J
I),
AI)w = (A+, + A+,)
substituting the value of AI), in above equation
Jw = L, (A$, + ... (11.7)
Thus, the inward or outward rate of flow or water from cell to cell and tissues can be
calculated from the above expression.
Let us now consider the flow of water in the plant through intercellular spaces
(apoplasm) where the limiting membranes are absent. Then the flux is given by
Jw = Hc AJIW
H, = hydraulic conductance
1
Because H, = -
R
R = Hydraulic resistance to flow of water

The water flux is directly proportional to AI) and inversely proportional to hydraulic
resistance (R). In other words, higher the A$, more will be flux but high R will
decrease the flux.
.
In plants water will move through the pathway which offers least resistance. Between
the two routes - cell walls and cell to cell, the membranes of the cells yxert more
resistance (because of low permeability) than the cell walls. Therefore, water can flow Hydrostatic pressure is the
relatively easily through Eel1 walls. Water will not experience the resistance of plasma Pressure exerted by or on aliquid
membrane when it moves from cell to cell via plasmodesmata. The xylem conduits above Or be'ow atmospheric
pressure.
which are not obstructed by cell membranes have least resistance and the rate of flow
of water is very high. The ratio of R in xylem, cell walls and cell membranes is in the
order of 0.3 : 1 : [Link] explains why xylem is the pathway for long distance
transport as has been observed experimentally. The resistance in xylem varies
inversely with the diameter of xylem elements. The smaller the diameter of xylem
greater will be the resistance.
In soil, pressure potential is insignificant and osmotic potential is zero because there
are no membranes (solute and water move together). Hence, the driving force AI),
in soil is determined by the matric pressure.
AJIw(Soil)= - AJlm(so,~)
The hydraulic resistance v ~ i i e sfrom soil to soil. The fine soil particles with small
space between them offer more resistance than coarse particles with large spaces.
When I), of soii falls R increases and then the plant take up less water.
Plant Physiology-1 Also, in the cell walls the driving force is determined by A$,. In thoie leaf cells
which are losing water rapidly, matric forces become significant and consequently JI,
of leaf decreases. So water moves from the wetter cells. Thus a continuous gradient
develops which operates along the cell walls throughout the plants. Of course, it will
break at points where cell walls are impregnated with hydrophobic substance for
example - Casparian strip.
SAQ 2
, . a) List the three factors that determine the value of $
, in plant.

..........................................................................................................
b) The water potential in a cup (JI,) containing salt solution will be
JIC ' JIW

JIC > 1IJw

JIC = $w

c) What will be the water potential of a plasmolysed cell if its osmotic pressure is
7.9 MPa.

d) Fill in the blank spaces in the following statements with appropriate words:
. i) At full turgor JI, of a cell will be .......................
ii) The net flow of water movement in a system will stop when $,,will be equal
and ...................... to JI,.
iii) Greater matric suction will ....................... the water potential in a system.
-- - -
1 1 7 GRADIENTS OF WATER ,POTENTIAL
"
The absorption of water by a plant involves the water relations of an individual cell,
a group of cells and finally of the whole plant. Therefore, we will consider water
relations at different levels of organisation. We have already discussed the long
distance transport of water.

Movemqnt of Water in a Single Cell

Isolated cells, single celled organisms and root hairs absorb water directly from their
surrounding media. Let us consider an ideal parenchymatous cell. The vacuole
occupies 90% of the cell and contains cell sap which is a dildte solution of salt and
other small molecules. Due to osmotic potential the cell sap has a lower water
potential than pure water. When such a cell is placed in pure water a gradient
develops due to the difference in water potential. This results in the movement of
water inside the cell (Fig. 11.12). However, in no time the concentration of sap also
decreases which lowers the osmotic potential of the sap. Thus, the difference between
the potential of pure water and cell sap gets reduced. This lowers the force by which
water enters the cell. We can represent the relationship with the following equation:
JIw (outside the cell) = ($, + $), inside the cell
where, +, is total water potential of the system, JI, is the osmotic or solute potential
and $,,is pressure potential due to cell wall pressure or turgor pressure. At full turgor
pressure the sum of JI, and JI, is zero. Hence JIw is zero.
The driving force F that causes water to move can be represented by the following
equation:
F = gradient (JI, - JIe)
where *,,is the total water potential of the cell including that of the cell sap and I),
is the total water potential of the external medium. If the latter is pure water then

*,
its value will be zero. In that case the driving force will be equal to I), However, as
the water will move into the cell I), will be regulated by and I),. In this relationship
the elasticity of the cell wall would also play an important role. The volume of the
cell would increase upto a certain limit with the dilution of the cell sap and this will
increase the total osmotic potential. This in turn would influence the force developed
due to the gradient between the cell and the medium.

1 Water flowing in from a

higher concentration of ,,
-
,
--- --- -- - - - -------------
,,- ' - - - - - - -
-, - - - --
-
water molecules to a lower '----
~ ~

concentration of water
-Water-
- --- ------ --
-- -- -- -- -
-------------------------------
- - -~

in the cell sap.

Flg. 11.12 :A single eeU surrounded by water.

In the cells of root, leaf and other parts of plant, the external medium is the water
in the cell walls and intercellular spaces (apoplast) which is under atmospheric
pressure and has very low osmotic potential. But the matric forces exerted by the cell
walls are higher, therefore, the water potential in apoplasm is determined by matric
forces exerted in ceil wall.

Water Relations of a T h u e
In higher plants no cell exists in isolation from others. Even [Link] hair which is
projected outside into the surrounding medium is attached with other cells on all the
remaining sides. In a transverse section, it would appear to be surrounded on the
three sides by other cells. Thus, the water relations of a root hair are governed on
one side by the surrounding medium and on the other by other cells. Let us just
consider two cells A and B joined with each other through a common cell wall. If
these two cells individually have the same total water potential, then there will not
be any net exchange of water between them. This can be shown as follows:
The total water potential of cell A ),*,( will be equal to the sum of its osmotic
potential and pressure potential (@,A + )*., Let us say that for the cells A and B
these values are
*WA = (*,A + *,A)
$WB + ($,B + $,B)

in the vacuole is lower than *, *,

In a cell the matric forces are much less. Here the is lowered because of +., If the
*, in apoplast, then water flows inwards.
Now, the driving force (F) will be the difference in the water potential of two cells.
= (*WA - *WB)
,+,
If is equal to JlwB the gradient will be zero and so will be the F. Therefore, no
net exchange of water pill take place between the cells A and B. However, we must
realise that the same value of, ,$ and,,$ doks not necessarily mean that the two
cells should have the same osmotic potential and the same turgor pressure or wall
pressure.
Plant I'hy siology-1 On the other hand, if the total water potential of cell B is lower than that of cell A,
a driving force will develop which would cause influx of water into cell B till the two
cells attain the same water potential. Now, this example can be extended to a larger
number of cells which are connected with each other in tissue. If there are 20 cells
beginning from 1to 20 they will attain an equilibrium amongst themselves, depending
i
on their total water potential in the same way as discussed for the two cells attached
to each other. Under such conditions a situation may come when water absorption
and movement will come to a standstill.
Water Relations of a Whole Plant
Tabk 11.1 : ~pproximatemagni- Let us now consider the water relations of a plant considering leaves, stem and roots,
tudes of wakr potentjPl io that provide a continuum for water in soil-plant-atmosphere system. The total water

i"
the soil-plan&atmosphere system potential in the atmosphere could be very low, depending upon the temperature and
Component Water Potential humidity. Table 11.1shows an approximate magnitude of water potential in the soil-
plant-atmosphere-system.
Soil -0.1 to -20.0
It is clear from the data in Table 11.1 that the difference in total water potential in
Atmosphere -100 to -2000
-5.0 to -50.0 the soil-plant-atmosphere system would generate a driving force for water movement
from the soil through the plant to atmosphere. If this continuum is broken, the driving '
force would automatically disappear.

SAQ 3
a) In a plant water moves from an organ A to an organ B bacause
i, $WA < $WB
ii) J l w ~> J l w ~
iii) Jlw, = $WB

b) Calculate the value of A+, if water potential of xylem is -0.5 MPa nt the base
of the tree trunk and -1.5 MPa at the top.

c) Water potential in a tree in three tissues A, B, apd C was found to be -0.4, -3.1
and -0.09 MPa. What would be the direction of movement of water.
.........................................................................................................

d) A raisin swells in water because

i) Jlw raisin > Jlw Or
ii) Jlw > Jlw raisin

11.8 WATER ABSORPTION

Water status of a plant is determined by two major factors: i) water absorption and
ii) water loss. We deal with water absorption in this section and water loss in the
following section.
Water absorption is regulated by soil factors, rate of transpiration and size and
distribution of roots. The soil factors that regulate water absorption are: i) soil water
content, ii) difference in water potential between soil and root, iii) concentration of
soil solution, iv) soil temperature and v) aeration of the soil.

Soil Characteristics
The physical properties of soil govern water-holding capacity and the water
availability to the plant. The fine soil particles of clay have much more water-holding
 Plant Water Relalions
capacity than silt and sand. Addition of humus increases the water-holding capacity
of soil. Water moves through pores present in soil. The pores form because individual
soil particles aggregate to form large particles of varying sizes called micelles. The
pores are spaces left between micelles. The size of pores can be small (micropores)
or large (macropores) depending upon the soil type. The pores get filled with water.
A soil freshly wet with irrigation or rain water cannot retain all the water. Much of
the water percolates through macropores due to gravity. This water has been termed
as gravitational water. The remaining water that is held tightly by hydrogen bonds to
the soil particles against gravitational forces is called capillary water. This water is
available to the roots. Part of the capillary water that is held very tightly and is not
available to the root is termed hygroscopic water.
The water content of a soil is expressed as

% of soil water = F W - D W x 100

DW
FW = Field weight of wet soil
DW = Dry weight (obtained by heating soil to 60°C)
Two terms field capacity (FC) and permanent wilting percentage (PWP) are often
used to describe the water status (capillary water) of the soil.
Field Capacity (FC)
It is the capacity of a field to hold the amount of water against gravitational forces.
It is expressed as the percentage of the dry weight of the soil. Field capacity
represents the upper limit of water availability and its value differs from soil to soil.
The capacity of clay soil on an average is about 40-45%, silt 20% and sand 5-10%.

Permanent Wilting Percentage (PWP)

It is the percentage of moisture in the soil at which a plant wilts and does not recover
unless water is added to the soil. PWP for clay is about 26%, for silt 10% and for
sand 3-5'10. It must be noted that PWP is used to express property of a soil not any
feature of the plant.
However, the moisture content at which any plant shows permanent wilting need not
be the same for all plants even in the same soil. This happens because some plants
are prone to wilting at fairly reasonable water content of soil while others wilt only
when the moisture content is very much reduced.
The water status of a plant in terms of FC and PWP is significant only if the soil
properties are known. This is because the soil determines the availability of water.
However, it is desirable as well as easier to express soil water status in terms of water
potential so that soils become uniform with respect to water.

Soil Temperature
Soil temperature is known to influence water absorption and ultimately transpiration
to a considerable 'extent. In many plants water absorption is reduced sharply below
10°C. Water absorption also slows down above 25°C. In most instances, temperatures
above 40°C in the rhizosphere does not support water absorption and plant may show
signs of wilting. The following are the reasons suggested for the reduced absorption
of water at low temperature: i) decreased root growth, ii) increased viscosity of
water, iii) increased resistance of water into roots due to decreased permeability
(increased hydraulic resistance) of cell membranes and iv) decreased metabolic
activity of root cells.

Soil Aeration and Flooding

It is not unusual to observe some plants wilting while they stand in water. The
following are the possible'reasons of flood injury.

i) Poor availability of 0, and accumulation of high concentration of CO, around

roots. .. .
ii) Changes in the pattern of ion uptake resulting in the accumulation of some ions
to toxic levels.
iii) Accumulation of toxic substances in the root and/or around them.
Plant Physiology-1 Poor availability of oxygen affects respiratory activity of roots, thus lowering the ATP
supply. In the following units you will learn that ATP is required for the active uptake
of ions. Low uptake lowers the osmotic potential of roots cells and hence water
cannot enter because A$, between root cells and soil is not sustained. Increased
concentration of C 0 2 affects the permeability of membranes and thus affects water
uptake adversely.

Among toxic substances are the metabolites produced as a result of anaerobic

respiration. These could be alcohol or aldehyde and even ethylene. Plants usually
become chlorotic under waterlogged conditions: This is apparently due to reduction
in translocation of iron from roots to the top of the plant.

Root
Root system is directly related to the absorption of water and its growth under field
conditions is very much influenced by soil. In dryland agriculture, particularly root
structure has apparently greater significance. The rates of water absorption into roots
of different plants differ in their stage of growth. Highest rates of water entry are
associated with root hair and unsuberised roots and lowest with suberised woody root.

Water Absorption and Transpiration

The rate of water absorption is controlled by the rate of transpiration. A high water
potential in the atmosphere would reduce water loss from the plant and consequently
slow down water absorption. But i t d r ~ snot fol!c.w from this that water will be
continuously absorbed if the water potential o i :he atmosphere is very low and soil
is near drying. When the equilibrium in the soil-plant-atmosphere system is disturbed
either due to soil or atmosphere, the plant can respond appropriately and can control
water absorption or water loses. We discuss below some of these aspects.

Let us first see the relationship of water ab'sorption and transpiration in a well
watered plant (Fig. 11.13). By and large all plants show a diurnal behaviour in their
rates of transpiration. There is a rapid increase in transpiration during the morning
hours and a peak is reached in the early afternoon. Then a decline begins and a
minimum is reached during the night.

------ Absorption

Fig. 11.13 :~ i u m a variations

l in rates of transpiration and absorption of water by a plant.

You may note in the graph that water absorption keeps pace with losses due to
transpiration except around noon when it lags behind. Why is it so? You have learnt
chat the resistance to the movement of water across the roots is generally higher than
in the leaves because in roots water moves through symplasm. In roots it faces high
resistance of [Link]'of cortical cell and that is why there is an absorption
lag. This idea is 'supported by an experiment where water absdrption was recorded.,
after removing the roots of a plant. As shown in the graph (Fig. 11.14) the absorption
lag is reduced greatly.
Let us now see what happens to water loss if'soil is not moist. Ralph 0. Slater (1967),
a plant physiologist at Australian National University, Canberra, studied the
 of Plant Water Relations
relationship between water potential of soil (4w,il), root (4, root) and leaf
a plant where water supply was withheld for five days. The graph (Fig. 11.15) shows
diurnal changes in leaf potential, water stress around the noon time and recovery of
lag in absorption at night. The water potential of the soil decreased almost linearly

Without roots
-3
% 0.10 - ------------
*-. /
#L..---

.a 0.08
8
- .4+4-e+4
4+ \-Shoot
\'\
covered
I #'
f
'
c--
With roots
:'\.,
8 -
#
-

$ 0.06 - f
5 .eeO

8 0.04 - q.
-*-+a
. ,J'
#
',

tShoot uncovered
4 0.02 -
0 ----I

230
. 10 20

Time/ Minutes
Fig. 11.14 :Canpmlpnof rated water absorption in rooted and rootless plant. W%entrurspimtlon Ls high
(rboot uneovacd), the rate of water abmrptlon In a rooted plant is less than rootiem plant.

after second day. It must be pointed out that when plants are under water stress,
water tends to move out of the cells that are not situated in the main pathway of
transport of water. This causes reduction in turgor, i.e. cell volume. In fact, it was
demonstrated q century ago by Josef Friedrich that there are daily fluctuations in the
diameter of tree trunk, shrinliage in the morning and recovery in the diameter by
night.

f
Y w leaf

30 d ~ i ~ h t
1 2 , 3 4 5 1 6
Days Day
Fig. 11.15 :Diagram ahowing m b l e changes in leaf water potentipl(gwU ) and root water p o b t i d
(Jlw d a tMlpIrins p h t rooted la sou allowed to dry from a water potential near zero to
a w8ter potfmtkl(gw at rbkh dtlng occmrs.

SAQ 4 --
Which among the foll'owing siggestions are desirable for growing healthy plants. ~ u i
a tick mark for the correct ones.
i) Excess use of fertilisers.
ii) Saturation of plants, particularly growing in pot, with water.
iii) High humidity area for water-sensitive plant,

iv) Use of very cold water in warm season.

v) Use of lukewarm water in cold season.
Plant Physiology-I
11.9 WATER LOSS
Plants lose about 98% of water to the atmosphere by transpiration. Often water loss
by transpiration exceeds gain by absorption and results in negative water balance
within the plant. Small and moderate deficit that occur due to high temperature
during the day are compensated during the night but prolonged deficit causes
iireversible damages and threatens the plant's survival.
Transpiration is essentially evaporation of water from the aerial portion of the plant.
However, evaporation of water from open surface meets less resistance while
evaporation of water from leaves faces considerable resistance. Transpiration occurs
mainly through stomata of the leaves. This is called ,stomata1 transpiration. About
5% of the water is lost from the leaf through the cuticle. This is called cuticular
transpiration. In woody plants there are lenticels opening within the bark that
. . these cells is called lenticular
function in gas exchange. The water loss through
transpiration.

11i9.1 Stomata /

The cross-section of a leaf shown in Fig. 11.16a shows the position of a typical stoma
(plural stomata) which however, differs from species to species, with respect to the
size of the pore, structure and size of the guard cells and depth and size of the
stomatal cavity. As indicated in the diagram b, water evaporates from wet mesophyll
cell walls that border intercellular spaces, the vapours then diffuse out through
sub-stomatal cavity and stomatal pores to the air outside the leaf. The water potential
gradient develops in the sub-stomatal cavity, stomatal pore, boundary layer and the
atmosphere. During transpiration the sub-stomatal cavity has relatively much higher
water potential as compared to the atmosphere, therefore, the water vapours move
out. This is turn lowers the water potential of the sub-stomata1,[Link],
the cells surrounding the sub-stomatal cavity evaporate water through their cell walls.
Depending upon'the water potential of the environment, the water potential of the
sub-stomatal cavity and the surrounding cells is lowered. This gradient eventually acts
as a 'pull' on the water column which maintains continuity through the vascular
bundles of the leaf. The intercellular spaces also play an important role in this respect
because they are in continuity with the sub-stomatal cavity and cause a gradient
quickly.

Upper epidermis
Stoma, with cuticle

Palisade
parenchyma

Spongy
parenchyma

Stornz~es Lower epidermis

with cuticle
(a) \
(b Stoma

Fig. 11.16 :a) Cross-section of a leaf showing the rplationship of stoma with other leaf tissues.
b) Stoma enlarged to show the path of water vapours from leaf to ahnosphere.

When water evaporates through leaves it experiences considerable amount of

resistance (Fig. 11.17) which can be grouped into the following two categories:
a) leaf resistance (internal resistance), and b) air boundary resistance (external
resistance).
 Plant Water Relation6
The components of leaf resistance are cuticle, mesophyll cells, intercellular spaces of
the leaf and stomata. The cuticular resistance is maximum followed by stomatal
resistance and air boundary layer resistance. The pathway of movement of water
vapours is analogous to the electric current moving in circuit, the greater the
resistance, the smaller the flow of water and vice versa.

Intercellular

Cuticle (r,)
Guard cell

Epidermal cell
~"bstomatalcavity rl

Fig. 11.17 :Scbematlc cross-seetion of a leaf showing different resistances.

The cuticle forms outermost surface of the leaf and offers resistance to the
evaporation of water vapour and entry of carbon dioxide necessary for
photosynthesis. Stomata perform the following main functions:
i) They allow entry of C 0 2 necessary for photosynthesis,
ii) They control water loss through transpiration and thus protect the plant from
desiccation,
iii) At higher temperature (above 35°C) they promote transpiration which serves to
cool the leaves.

Stomatal resistance is most important because gas exchange between leaves and
external atmosphere takes place entirely through stomatal pores. Stomatal resistance
depends mainly on the size and shape of the stomatal cavity and size of stomatal
aperture. Can you tell which pore size, small or large would exert greater resistance?
Of course, smaller the pore greater would be the resistance for the outward
movement of water vapour. On an average a standard pore measures about 20 pm
in length and about 11 pm wide at its widest point when fully open.

Water loss from the leaf also depends on the number of stomata per leaf. One of the
aims of agricultural scientists is to find ways of minimising the water loss from the
plants so as to increase the efficiency of water use. One way is through a study of the
number of stomata per leaf. Some investigations have been carried out keeping the
following questions in view:
i) What is the distribution of stomata and how is it related to water loss and carbon
dioxide intake?
ii) How can water loss be reduced without seriously impairing carbon dioxide
intake?
iii) Is it possible to evolve varieties which possess characteristics associated with low
evaporation of water?
There are two parameters normally used for expressing the distribution of stomata.
a) Stomatal frequency : The number of stomata per unit area.
b) Stomatal index :The ratio of number of stomata to the total number of cells per
unit area.
Plants differ in their stomatal frequency. In monocots the upper and lower surfaces
usually have the same frequency but in dicots the lower surface usually has a higher
frequency than the upper surface. However, the frequency of stomata can change
with the position of leaves on the plant.
 Studies have been carried out to see if any relationship exists between i) stomatal
frequency or stomatal index and transpiration and ii) stomatal opening or stomatal
index and photosynthesis. Using two barley lines which were high and low in stomatal
frequency Misken and coworkers (1972) found that stomatal resistance and
transpiration rates differed significantly between lines but photosynthetic rates were
the same. The lines having low stomatal frequency had higher stomatal resistance,
transpired less water than lines having more stomata. A decrease of 25% in stomatal
frequency reduced transpiration rates by about 25%. However, the photosynthetic
rate was unaffected by decrease in stomatal frequency. These studies suggested that
it should be possible to alter transpiration withodt altering photosynthesis by selecting
varieties with fewer stomata in barley o f other crops.
Let us first have a close look at the stomatal aperture. Fig. 11.18 illustrates an open
(a) and a closed (b)' stoma in surface view. As you may know each stomatal pore is
surrounded by a pair of guard cells. In closed stoma guard cells appear like two joined
kidney beans. In monocots guard cells are dumbell shaped and are arranged in pairs
in contact at the bulbous ends (Fig. 11.1&). In some species the epidermal cell
adjoining the guard cells are specialised and are called subsidiary cells (Fig. 11.18d).
It is important to mention here that guard cells contain chloroplasts and mitochondria
and can synthesise starch while no other epidermal cells contain .cl~loroplasts.

11.9.2 The Mechanism of Stomatal Opening

It has been known for over a century that stomata open because of reversible turgor
changes in the guard cells. Stomata open when turgor in the guard cells is high and
close when it is low. It is clear that the turgor would increase only if the solute contelit
of guard cells would be higher than the neighbouring epidermal cells. Falling of solute
content would decrease turgor.
How do the two guard cells on swelling form an aperture? The cell wall of guard cells
is peculiar because the cellulose microfibrils which constitute the major stnictural
feature of plant cell are arranged radially extending from the centre towards the
periphery (Fig. 11.19).

Closed \ /'
Cellulose microfibril
Opn

I
Own
cell
Epidermal a l l

Fig. 11.18 : Surface view of an This arrangement restricts the expansion of cell wall in transverse directio?. Since the
open a) and a closed b) stornatal inner wall (towards the pore) is thickened and is less elastic than the outer wall, the
aperture in a leaf of a dicotyledon uneven longitudinal expansion causes the cells to arch away from each other forming
C) and d) of momcotyledon a pore in the centre.
e) guard cells surrounded by
subsidiary cells. Before we discuss the mechanism of control of stomata, it is important to bear in.
mind the following observations made in this regard.
i) Normally, stomata are open in the day and are closed at night. However, a drop
in supply of water leads tp the day time closure of stomata.
ii) Stomata open when the internal concentration of C 0 2 drops and close when the
internal concentration of C 0 2 is maximum.
iii) ark C 0 2 fixation (CAM plants, refer to Unit 13) occurs in-the guard cells.
 I'lar~tWater Relations
iv) In guard cells there is a change of pH in light (day) and dark (night) which is
associated with the interconversion of starch and sugar.
v) Opening and closing of stomata are related to the osmotic potential of the guard
cells and the permeability of membranes.
vi) At the time of opening of stomata, there is an inflow of potassium ions into the
guard cells but when the stomata close, the guard cells lose their acquired
potassium to the surrounding cells.
vii) Inhibitors of cyclic phosphorylation can also close stomata.
viii) Blue light also brings about changes in stomata1 movement - serving to open
stomata.
ix) Abscisic acid - a plant hormone, at a very low concentration can lead to the
closure of stomata.
There are two ways in which the relative turgor of guard cells may be altered. i) a
decrease in osmotic potential or ii) a decrease in pressure potential. In both instances
the water potential of guard cells will fall and hence water from neighbouring cells
would move in. Alternatively, guard cells may face mechanical pressure from the
subsidiary cells if they face a sudden change in their turgor.
There is overwhelming evidence that the osmotic potential of the guard cells
decreases due to the migration of K + ions from the surrounding cells into the guard
cells (Fig. 11.20). It is also observed that the uptake of K + requires ATP which may
be generated by degradation of starch through respiratory metabolism and
photophosphorylation. ATP is utilised to pump out protons out of the guard cells by
a membrane bound ATPase (refer to section 8.4 LSE-01). The supply of protons is
maintained by malic acid which is synthesised from stored starch. Due to the active
pumping of protons, an electrochemical gradient develops across guard cell
membrane. Now the ions can move passively into the guard cells. In some species
C1- ions or other anions move along with K+ in and out of the guard cells. Thus,
there is a decrease in osmotic potential due to accumulation of K + , C1- and/or
potassium salt of organic acid mostly malate. Consequently, water enters guard cells
and builds up turgor. Reverse events would bring about closure of stomata.

Malate -----

Fig. 11.20 :Changes in the turgor of guard cells by the movement of K+ and other ions. Solid lines indicate
active transport, broken line passive fluxes and? mark uncertainty. Upward arrow indicates
rising level of malete (e) and downward arrow falling level (b).
Plant Physiology-1
11.10 FACTORS CONTROLLING STOMATAL
APERTURE
'Khul Ja Sim Sim' the two rocks slide and the gates to a great treasure open. With
these magic words "Ali Baba and Chalis Chor" could open the door. Let us now find
out what is 'Khul Ja Sim Sim' for moving the guard cells of the stomata, so that C 0 2
can enter the stomata and make food for organisms of the earth. You have learnt
that more than one factor can trigger the change in the turgor of the guard cells and
bring about stomatal movements and control the size of stomatal aperture. The main
controlling factors are:
i) light,
ii) the level of C 0 2 , and
iii) high temperature.
To understand better you can imagine the factors as hands that can open and close
stomata by moving the turgor operated valve - the guard cells (Fig. 11.21). The
stomata open when the level of C 0 2 falls due to its rapid utilisation in photosynthesis
during the day. However, it is not known how guard cells sense the low level of C 0 2 .
The highest rates of photosynthesis are obtained when the conditions for light and
temperature are optimum and plants are well watered. But if the water potential in
the leaf falls under low level of C 0 2 , then conserving water b e c ~ m e smorc Lrgent
than food production for the plant and the stomata close. In other words, the water
status of plant overrides the control by C 0 2 .

Turgor-operated
valve
Conserving water feedback

CO2
Temperature feedback (above 350C)
7
(T7 - (
'
feed back

I '-7' C02 sensor

' &IT<
temperature C02 sink Tissue water

Fig. 11.21 : A simplified model showing the etYect of various factors on stomatal aperture.

The rate of photosynthesis increases with increase of temperature but above 35°C
cooling of the plant becomes necessary. Therefore, irrespective of C 0 2 concentration
stomata open widely above 359C provided that there is no shortage of water. Here
again the temperature overrides the control by C 0 2 concentration. This makes sense
because high temperatures are deleterious to the health of the leaves, and
evaporation-of water through transpiration can lower the temperature of leaves.
Before we close this unit, it is important to point out that the water loss by a plant
due to transpiration is very high usually in the range of 0.1 to 2.5 g dm-2h-' during
day time. However, it plays no major role in growth and development of plant so far
is known. We have mentioned above that it cools the plant at high temperatures.
Perhaps, the transpiration stream facilitates the availability of mineral ions to all parts
of the plant. Soil contains minerals in very dilute concentration but in cells they get
accumulated in much higher concentration due to active transport aided by their rapid
availability in the transpiration stream.

You have learnt that the external application of plant hormone abscisic acid (ABA)
to leaves causes stomata to close. During water stress, increase in the level of ABA
of the leaves has also been observed. Therefore, it is quite likely that ABA is an
internal control for the regulation of water content. Phenyl mercuric acetate - a
fungicide has been used as foliar spray in low concentration ( ~ o - ~ M It
) . brings about
partial closure of stomata for about two weeks without visible damaging effects to
the plant. Other chemicals such as colourless plastics, silicon oils and low-viscosity Plant Water Relations
waxes are also used as foliar spray. These chemicals form on leaves a film permeable
,
to C 0 2 and O2 but not to H 2 0 . Water-saving strategy by reducing transpiration
though seems quite promising, yet it is a big challenge for plant physiologists.

SAQ 5

a) List the features of guard cells essential for opening the stomata.
'

11.11 SUMMARY
Water is the key molecule for the maintenance of life on the earth. About 85 to
90%' of a plant is made up of water. The quantities of water used by plants are
enormous.
Water is a good solvent and reactant in the cell. For plants,water is also crucial
because the hydrostatic pressure of water provides turgidity to the cells, so that
the soft stem parts are able t o stand erect.
Water moves up into the trees due to negative hydrostatic pressure created in
xylem by the transpiring leaves. The continuity of water column in the xylem is
maintained because of cahesive property of water.
In plant tissue water molecules move through two routes -
apoplasm and
symplasm. Apoplast transport is through non-living portion of the plant and
symplast is through cell to cell via plasmodesmata.
Water status in a plant is expressed by water potential ($,). The rate of flow and
the direction of movement depend upon the gradient of water potential between
the two points. Water tends to move from a region of high water potential to a
region of low water potential.
Water potential is affected by solute concentration, pressure and absorptive or
matric forces. Addition of solute lowers the water potential while increased
pressure' on water causes a rise in water potential. Absorptive forces between solid-
liquid interfaces lower the water potential. Water potential is computed as the
algebraic sum of component forces: osmotic potential (due to solute), pressure
potential (due to hydrostatic pressure, matric o r suction potential (due to adhesive
forces).
In a cell both solute concentration and turgor pressure can have offsetting effect
on water potential. The combined effect of the two in cells can build up the
gradient and determine the rate of flow and direction of water movement.
The gradient of water potential -the driving force for water movement in soil is
due to differences in matric potential. This gradient also occurs along the cell walls.
Water movement within the dead xylem cells occurs along the gradient of
hydrostatic pressure.
There is resistance to water flow in the soil and in the plant. Water takes up the
path of least resistance. Membranes of cells exert greatest resistance while xylem
offers least resistance.
Water status of a plant depends upon water absorption and water loss due to
transpiration. Soil characteristics, soil temperature, aeration, flooding and
"
structure of roots are the factors which affect water absorption.
The rate of water movement in plant is determined by the rate of transpiration in
a well watered plant. Temperature, wind velocity, humidity of the air, light
intensity and degree of stomata1 opening affect the rate of transpiration. However,
ultimate limit to transpiration is the supply of water from soil.
 Among stomatal, cuticular and lenticular transpiration, stomatal transpiration
accounts for major water loss. Evaporation of water faces considerable resistance
from the leaf - cuticle, mesophyll cell, intercellular spaces, stomata and air
boundary layer of the leaf.
Stomata control water loss through transpiration, allow gas exchange and control
temperature of the leaf. Some efforts have been made to reduce water loss through
stomata without affecting C 0 2 intake.
Stomatal aperture is controlled by the changes in the turgor of guard cells. The
turgor in turn is affected by the metabolism and the movement of K+ and other
ions into and out of the guard cells. A proton pump powered by ATPase operates
in the plasma membrane of the guard cell and the K+ uptake is a passive uptake.
Stomatal aperture is controlled by the level of C02, light and high temperature.
However, if conserving water or saving plants from deleterious effects of high
temperature becomes more urgent than food production then these factors
override the system.

- - - -
11.12 TERMINAL OUESTIONS
1) Why do animals need less water than plants?
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2) What would happen if a twig of a potted plant is cut into two pieces and joined
with tygon tubing in such a way that it leaves an air gap between the cut ends?
Will water continue to move?

3) A plant cell with an osmotic potential of -7.6 bars and pressure potential of 3
bars is placed in a slide with a drop of pure water, in which direction the water
will flow? What is the $, $p and $, of the cell and water initially and after
equilibrium?

4) Draw the structure of stomata and explain how stomatal aperture opens by the
increase in the turgor of the guard cells.
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11.13 ANSWERS
Self-assessment Questions
1) i ) F , ii)F, iii)T, iv)T
 Plant Water Relations
2) a) Osmotic potential (JI,), pressure potential (JI,) and matric potential (JI,)
b) JI, = JI,. The JI, of salt solution in a cup will be zero because there are no
membranes present. Therefore, its water potential is equal to pure water.
c) -7.9 MPa.
We know that ?r = -+,. Therefore, the osmotic potential will be -7.9 MPa.
Since JIp = 0, therefore, JI,, = JI,
d) i) zero ii) opposite iii) lower

b, = kv Base - *w top
= -0.5 - (-15)
= -0.5 1.5 +
= 1 MPa
c) A = -0.4 MPa, B = -3.1 MPa, C = 0.09 MPa
JIw= 0.09 > -0.4 > -3.1
the flow of water will be from C + A -,B
d) ii)
4) iii) and iv) are the right suggestions
5) a) i) A thickened cell wall, facing the pore,
ii) presence of chloroplasts,
iii) supply of ATP,
iv) a proton pump in the membrane,
v) the capacity to generate higher turgor pressure than adjacent cells.
b) i) F, ii) T, iii) T, iv) F.
Terminal Questions
1) In animals a great deal of water is recycled through the body in the form of blood
plasma and other fluids in vertebrates. In plants more than 90% of the water
absorbed by lkaves is lost into the air as water vapour through the process of
transpiration.
2) The plant will absorb water but the uppeipart of the plant will wilt. This is .
because the air column between the two stem pieces does not provide a
continuum. In plants the movement of water takes place only in liquid phase
which can stand enormous pressure without breaking the liquid column.
3) Water will flow into the cell until the water potential of cell and water drop is
equal.
The following are initial and final (equilibrium potentials
Initial Equilibrium
water drop cell water drop cell
$7, 0 -7.9 0 -7.9
+P 0 +3 0 +7.9
3rw 0 -4.9 0 0
4) Please see text for the answer.