Unit 1 Plant - Water Relation

UNIT 1
PLANT –WATER
RELATIONS

Structure
1.1 Introduction Osmotic Pressure

Objectives Diffusion Pressure Deficit

(DPD)
1.2 Life Supporting
Characteristics and Chemical Potential
Biological Importance of Components of Water
Water to Plants Potential
1.3 Water Transport Gradients of Water Potential
Diffusion 1.4 Summary
Imbibition 1.5 Terminal Questions
Osmosis 1.6 Answers
Membrane Permeability

1.1 INTRODUCTION
Water is the most important single component which supports all living
organisms on earth. All animals and plants, whether terrestrial or aquatic, are
dependent on water for their survival. We could survive for some days without
food, but it is impossible to live without water.

Our daily weight fluctuates on account of excess or loss of water. Water is the
most abundant component in any living system. It constitutes nearly 70% of
the total body weight.

In plant cells most of the water is localized in the vacuoles (upto 95%) and the
rest resides in the peripheral cytoplasm. The amount of water in plant varies
among different species depending upon their structure. Aquatic plants such
as Chlamydomonas, Spirogyra, Chara, red algae, and others contain 97-98%
of their weight as water. This is true also for Azolla, Eichhornia and other
freshwater plants. However, among terrestrial plants, whether they are crops
such as wheat, rice, maize, or trees such as sheesham, mango, neem and 9
Block 1 Water and Mineral Nutrition
many 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 during the growth of an organ such as leaf or
seed. In this unit we will discuss the properties and importance of water for
plants.

Objectives
Objectives
After studying this unit, you should be able to:

explain the unique properties of water which make it the elixir of life;

differentiate between the processes of diffusion, osmosis and imbibition;

explain the various components of water potential and the relationship

between them; and

describe and relate the movement of water in a single cell to a tissue

and to the whole plant.

1.2 LIFE SUPPORTING CHARACTERISTICS AND

BIOLOGICAL IMPORTANCE OF WATER TO
PLANTS
The life-supporting characteristics of water are due to its unique physical and
chemical properties. Nearly 75% of our planet is covered with water in the
form of oceans and other water bodies. The dominating presence of water in
most cells must be seen in terms of its much larger number of molecules as
compared to others. This is because water has a low molecular mass (18
Daltons).

Water as a Chemical Reactant

Water in a liquid state is an ideal medium facilitating the mobility of different

macro/micro molecules within and across the cells. Water also influences the
structure of different chemical components of cells. Water participates in many
biochemical reactions either as a reactant itself (e.g., photosynthesis) or a
product in others (e.g., respiration). During photosynthesis water splits in light
reaction (photolysis) into oxygen and hydrogen. The latter is used in carbon
fixation through CO2 for making glucose and constitutes the basic reaction of
the autotrophic food chain. Various life characterizing biochemical reactions
like condensation, hydrolysis, oxidation, and reduction occur in an aqueous
medium.

Water for Seed Germination

Seeds do not germinate unless they get enough water to hydrate their
protoplasm. Hydration triggers various metabolic reactions, and the
10 metabolites are mobilized in an aqueous medium.
Unit 1 Plant - Water Relation
Water as a Solvent

Water is a polar molecule and has exceptionally high hydrogen bonding ability.
This unique property allows it to dissolve a variety of molecules like sugars,
amino acids, carrier proteins like hemoglobin and myoglobin. Sugars and
proteins which contain polar –OH or –NH2 groups easily dissolve in water.
Thus, an aqueous medium would be extremely/ highly suitable for uptake of
dissolved mineral nutrients-so vital for growth and development of plants. It
was precisely this property of water which made possible a variety of reactions
in the “hot primeval soup” of the oceans during the origin of life.

Thermal Properties of Water

Water is unique as it remains in liquid form over a wide range of temperatures

where life exists comfortably. If one considers the molecular size alone, water
must be expected to exist in a gaseous state in most of the areas on earth.
However, this is not true. The melting and boiling points of water are much
higher than what would be theoretically expected. Presence of oxygen has
introduced polarity and the tendency to form hydrogen bonds. It is this
extensive hydrogen bonding which gives water its unique thermal capacity i.e.,
high specific heat, thermal conductivity, latent heat of vaporization and heat of
fusion.

Adhesion and Cohesion

Adhesion and cohesion essentially refer to the “stickiness” that water

molecules have for each other and for other substances. These attributes are
due to the extensive hydrogen bonding. Water drop remains a drop on
account of cohesion i.e., mutual attraction between molecules. For this to
occur, the area of an air-water interface needs to be increased. This would
require breaking of the hydrogen bonds and energy input. This energy needed
to increase the surface area of a gas-liquid interface is called surface
tension. High surface tension makes the water drops spherical and help
movement of capillary water through the micro spaces. Another related
characteristic is adhesion, which is the attraction of water to a solid phase. In
plants,water “sticks” to the cell wall of the tracheary elements, i.e., xylem. This
again is on account of extensive hydrogen bonding. Both adhesion and
cohesion contribute to another related property, i.e., tensile strength. Tensile
strength can be defined as maximum force per unit area that a continuous
water column can withstand before breaking. This property is characteristic of
metals. Water is an exception and exhibits a high tensile strength of 30
megapascals (MPa).

The trio of adhesion, cohesion and surface tension gives water another unique
property of capillarity, i.e., the ability of a water column to rise as a
continuous column in plants.

Providing Turgidity to the Cells

Uptake of water results in turgor pressure which inflates the semi-elastic

cellulosic cell wall resulting in initial volume expansion. Proteins,
carbohydrates along with other metabolites are deposited only later during
growth. 11
Block 1 Water and Mineral Nutrition

SAQ 1
Match the terms in Column A with those of Column B

Column A Column B

a) Temperature regulation 1. Dipoles

b) Mutual attraction 2. 0.239 calorie

c) Hydrogen bond 3. Heat of vaporization

d) Orientation towards (+ve) 4. 10 picoseconds

and (– ve) ions
e) Hydration 5. Germination

f) Joule (J) 6. Cohesion

1.3 WATER TRANSPORT

How does water flow in and out of the cells? How is water taken up from the
soil by the roots? How does water escape from the plant into the atmosphere?
To answer these questions, you would need to understand the dynamics of
water flow with reference to the movement of substances and the different
forces which are operative in this movement. Particles also exert forces by
themselves. Before you study about them in detail, you must know that they
are diffusion, imbibition, mass flow, and osmosis. Translocation is the term
which broadly defines the movement of substances from one region to the
other. The process of translocation may be passive on active, depending upon
the need for expenditure of energy.

The movement of water occurs through cell to cell by two principal pathways:

a) Symplastic pathway where water moves from cell to cell via

plasmodesmata. Since cells are joined via intercellular connections, the
entire living portion of the plant forms a continuous single entity and is
called symplasm.

b) Apoplastic pathway is another route for the transport of water through

cell walls, intercellular spaces, and non-living cells of the xylem. The
non-living portion of plants is called apoplasm.

1.3.1 Diffusion
You can smell from a distance the aroma of food being cooked in the kitchen.
A bottle of perfume opened in one corner can be smelt in another corner. A
crystal of copper sulfate dissolves in a beaker of water turning the latter blue.
So does a drop of blue ink dropped in water. All the above phenomenon
highlight a common point i.e., the random movement of particles is caused by
12 the inherent kinetic energy they possess.
Unit 1 Plant - Water Relation
The particles strike one another repeatedly thus, resulting in their dispersal in
different directions. Also, these particles would always tend to move
(perpetually move) in the direction where there is minimum obstruction offered
by the other particles. It would mean, therefore, that the net movement would
be favored from the area of higher concentration to the area of low
concentration (region of higher free energy to a region of lower free energy).
Thus, the spontaneous process or the tendency of particles (ions, atoms, or
molecules) of gases, liquids and solids to get evenly distributed through the
available space due to their random kinetic motion is called Diffusion (Latin
diffuses = spread out). In other words, diffusion is a spontaneous process that
leads to directed net movement of a substance from a region of higher
concentration to a region of lower concentration, or simply as the net
movement of molecules from a region of high free energy to a region of low
free energy (see Fig 1.1). The term was coined by R. Brown in 1928.

After sometime the particles/molecules will evenly spread throughout the

available space. This would mean an equilibrium where the movement of
particles will be equal in all directions, and thus net movement will be zero.
The process of diffusion can be explained based on Fick’s First Law (a
quantitative description of the process formulated by Adolf Fick in 1855), which
in its oversimplified way suggests that the rate of diffusion is directly
proportional to the cross-sectional area of the diffusion path and to the
concentration vapor pressure gradient. At the same time rate of diffusion is
inversely proportional to the length of the diffusion path.

(a)

(b)

Fig.1.1: a) Diffusion of solute molecules from a region of high concentration

(A) to a region of low concentration (B) till the molecules are uniformly
distributed; b) At the time of uniform distribion, there is no net duffusion.

It is believed that diffusion is most effective over shorter distances, as the

average time required for a substance to diffuse for a given distance increases
as the square of that distance. By this analogy, it is easy for glucose to diffuse
in water, and with a diffusion coefficient of about 10-9 m2 s-1, a glucose
molecule can diffuse across a cell of about 50/µm diameter in just 2.5 13
Block 1 Water and Mineral Nutrition
seconds. Interestingly, the time taken by the same glucose molecule to diffuse
to a distance of 1m in water will be nearly 32 years !!! Thus, diffusion is not
much effective for translocation over long distances (see Unit 8).

The ions particles/molecules which diffuse through the medium are the
An example of solid
to solid diffusion is diffusate while the medium into which these particles disperse is the
when aluminum diffusant. For example, perfume is the diffusate and air is the diffusant; sugar
diffuses into silver is diffusate and water is the diffusant. Diffusion can take place between gas-
when both are placed gas, gas-liquid, liquid-liquid, solid-liquid and solid-gas and solid to solid
in contact.
phases. Rate of diffusion depends upon several factors.

a) Temperature: Diffusion is directly proportional to temperature as it

increases the kinetic energy of diffusate particles. Diffusion show a Q10
of 1.2-1.3. (The Q₁₀ denotes temperature coefficient, which is a measure
of the rate of change of any biological or chemical system because of
increasing the temperature by 10° C).

b) Density of diffusate: Diffusion rate is inversely related to the square

root of the density of the diffusate (d).

D∝
√

D = Diffusion rate,

d = density of diffusate.

Thus, gases diffuse more readily than do the liquids while the solids
diffuse the slowest.

c) The pressure exerted due to the tendency of the diffusate to diffuse is

called its diffusion pressure.Because of their inherent kinetic energy,
the movement of molecules will take place from the region of higher
diffusion pressure to region of their lower diffusion pressure.

d) Density and concentration of diffusant also determines/influences the

rate of diffusion. More concentrated and dense the diffusant,
lesser/slower will be the rate of diffusion.

Importance of Diffusion

The principle of diffusion is of great importance in plants as it is involved in the

movement of almost all substances. Processes such as uptake of water and
minerals, osmosis, transpiration, exchange of gases during photosynthesis
and respiration, short distance translocation in the symplast, spreading of ions
and other substances throughout the protoplast and wetting of cell walls, and
release of odor for pollination depend in part on diffusion.

1.3.2 Imbibition
Imbibition (Latin imbibere : to drink) is a process that involves absorption of
water or any liquid by the solid particles of an absorbent substance without
forming any solution. The liquid or water which is imbibed is called an
14 imbibate while the insoluble substance that imbibes water/substance is the
Unit 1 Plant - Water Relation
imbibant. The imbibants are usually in the form of colloidal particles which
hold liquid water adsorbed on their surface as well as inside the minute spaces
between the particles themselves. Although the liquid involved in imbibition in
plants is water, other substance may also act as imbibants. For example,
rubber imbibes ether/kerosene oil rather than water. Most important among
the plant imbibants are the hydrophilic colloids like proteins, pectins, starches
and cellulose. These substances differ in their imbibition capacities. The
phycocolloid agar-agar has the maximum imbibition capacity and can imbibe
almost 99 times its weight of water. Proteins also have a high imbibition
capacity of imbibing nearly 15 times water their own volume. Cellulose and
starch are poor imbibants and lignin does not imbibe water.

Agar agar > oily seeds > proteinaceous seeds > starchy seeds.

You must have observed at home that gram seeds when soaked, swell much
more than those of rice or wheat.

Since imbibition holds water in between the solid particles, the immobilized
water loses its kinetic energy. The lost kinetic energy is released in the form of
heat, often termed as heat of wetting. Kneading of wheat flour results in an
increase in temperature. Moreover, imbibition results in increase in the volume
(swelling) of the imbibant. However, this increase in volume is less than the
volume of water imbibed.

The process of imbibition develops a high pressure called imbibition

pressure. It can be defined as the maximum pressure which can develop in
an imbibant when brought in contact with water without allowing the imbibant
to swell up.

You will read in the following subsections that imbibition pressure is also
referred to as matric potential. Dry seeds show a matric potential value of
nearly –100 bars. Dry seeds like pea on encountering water can develop high
imbibition pressure of up to 1000 atm (other equivalents used are bars and
kg/cm2). 1 bar=0.987 atm=106 dynes/cm2=1.019 kg/cm2=14.5 lb./in2.

This property has been exploited since ancient times to break rocks by filling
dry wood in natural grooves of rocks and watering them. The wood swells up
and cracks the rocks.

Various factors influence imbibition:

a) Rise in temperature enhances the rate of imbibitions.

b) Texture of the imbibant, i.e., cohesion of molecules is inversely

proportional to imbibition capacity. So, gelatin, with loosely packed
particles will imbibe much more than hard wood.

c) Since the imbibants are predominantly colloidal, imbibition is influenced

directly by pH of the medium. For example, a negatively charged colloid
like cellulose will imbibe maximum at alkaline pH. Since proteins are
amphoteric, their absorption is maximum in other alkaline or acidic
medium, but least in neutral medium. 15
Block 1 Water and Mineral Nutrition
d) Electrolytes are imbibed at slower rates as they tend to naturalize the
charges of the imbibants.

e) Seed germination primarily depends on imbibition of water.

f) Imbibition results in the breakage of seed coat.

g) Many dehiscent fruits like pods of legumes, cotton bolls dehisce through
imbibition in dry conditions by xerochasic, hygroscopic movements.
Elaters of bryophytes also show hygroscopic movements.

1.3.3 Osmosis
The term osmosis (Gr Osmos = impulse) was coined by Nollet in 1748 who
first observed that a cylinder full of wine whose mouth was covered with
animal bladder started swelling up and even burst when placed in water.

Osmosis can be defined as the diffusion of water (or solvent) from a dilute
solution into a stronger solution when separated by a semipermeable
membrane. Thus, during the process of osmosis the direction and rate of flow
depends on a semi permeable membrane, concentration difference and
pressure difference in the two compartments.

Osmosis can be demonstrated by a simple thistle funnel experiment which you

must have performed in your schools.

A solution which can cause entry of water into it is called an osmotically

active solution. The osmotic entry of water into a system, organ or a cell from
outside is called endosmosis, while exosmosis is the withdrawal of water
from a cell or an osmotic system in response to a hypertonic solution. Swelling
of water soaked dry raisins with intact stalks is an example of endosmosis
while the fresh grapes shrink in size and become wrinkled due to exosmosis,
when placed in 15% salt or sugar solution.

Three types of solutions can be classified, based on the cell sap.

a) Hypertonic solution : A solution having more concentration than that of

the cell sap. If a cell is placed in such a hypertonic solution, water will
diffuse out due to exosmosis resulting in shrinkage of the protoplasm
(plasmolysis).

b) Hypotonic solution : In this case, the concentration of the external

solution is less than that of the cell sap. When a cell is immersed in a
hypotonic solution, water diffuses into the cell (endosmosis) resulting in
an increase in cell volume.

c) Isotonic solution : When the cell sap and the external solution have the
same concentration. There is no change in the cell volume as there is no
net movement.

1.3.4 Membrane Permeability

The tendency or ability of a membrane to allow or restrict the passage of
substances through it is referred to as Membrane Permeability. This is a
16 characteristic feature of all membranes. Rate at which the passage of solute
Unit 1 Plant - Water Relation
particles is allowed through the membranes varies with the permeability
property of membranes. For example, a permeable membrane allows the
passage of both the solute as well as the solvent particles through it. Majority
of cellulosic cell walls behave as permeable membranes. On the other hand,
the suberized walls of cork cells, or cutinized walls do not allow the passage of
any substance and are, therefore, impermeable. Semipermeable are those
membranes which allow only the solvent but not the solute particles to pass
through e.g., parchment, copper ferrocyanide membrane or collodion
membrane. Plasma membrane, tonoplast and membranes of organelles have
unique permeability characteristics. These differentially permeable or
selectively permeable membranes allow the passage of solvent (water) as
well as selected (specific) solutes by a variety of mechanisms, operative within
their lipoprotein complexes. For example, special protein-lined channels called
aquaporins are responsible for water absorption and its further packaging. In
addition, ions of solutes are transported through special ion channels
(approximately 30 kDa).

1.3.5 Osmotic Pressure

It is the hydrostatic pressure developed in a solution which is just necessary to
prevent the flow of water or a pure solvent to a solution (osmosis) when the
two are separated by a semi-permeable membrane. In other words, it is the
maximum pressure which can develop by an osmotically active solution when
it is separated from its pure solvent by a semi permeable membrane.

Osmotic pressure (OP) can be measured by an osmometer, which involves a

manometer. Pfeffer’s osmometer and Hartley’s osmometer are the two most
used instruments for this purpose. The former employs very dilute solutions
whereas the latter set up uses an external pressure source to counter the
osmotic pressure. Since both the entities are equal, OP can also be alternately
defined as the pressure that is required to be applied to a solution in order to
prevent an increase in its volume due to the inherent tendency of water (on
solvent) to enter it, when the two are separated by a semi-permeable
membrane.

The value of OP is directly proportional tothe concentration of the solution. The

highest value of OP is observed in halophytes (>200 atm). Osmotic pressure is
denoted by the following equation given by van‘t Hoff (1887).

OP = m RT
or
OP = c RT
Where OP = osmotic pressure
m/c = molar concentration of solution (concentration of solute/litre)
R = Universal gas constant (0.082 atm/mol)
T = Absolute temperature in Kelvin (K) = (t + 273)
t = temperature in °C
Theoretically, OP of one mole glucose solution at 0°C will be
OP = 1 × 0.082 × 273 17
Block 1 Water and Mineral Nutrition
22.38 = # 22.4 atm, = 22.69 bars (1)

= 2.269 MPa = 22.69 ×105 Pa,

and at 20°C will be

= 1 × 0.082 × (273+20)

= 1 × 0.082 × 293 (2)

= 24 atm = 24.31 bars = 2.431 MPa

= 24.31 ×105 Pa

The above-mentioned formula for OP is applicable to non-electrolytes e.g.,

glucose and sucrose. For electrolytes like NaCl or KCl, the degree of
dissociation will also have to be considered.

For electrolytes, the formula becomes OP = ciRT

Where i is the van ‘t Hoff’s ionization constant. Ionization increases the

number of particles in a solution. Thus, it is natural that the same volume, of
electrolyte would exert more OP than its equimolar non-electrolyte counterpart
at the same temperature and pressure. To give you an example, 0.01 M
glucose (non-electrolyte) will have an OP of 0.24 atm whereas equimolar
sodium chloride (electrolyte) will have an OP of about 0.47 atm.

Thus, the OP of an electrolyte is greater than that of the equimolar non-

electrolyte. Therefore, water moves from lower OP towards the higher OP.

Turgor Pressure (Hydrostatic Pressure/Pressure Potential)

The pressure developed by the turgidity in a cell is called turgor pressure

(TP). Turgor pressure is applicable to living cell and not to a free solution. TP
is called as hydrostatic pressure or pressure potential. When a cell is
immersed in water, water enters the cell as cell sap has a higher osmotic
pressure. It causes swelling of the osmotic system. As a result, the hydrated
cytoplasm presses against the cell wall.

The outward force applied by the turgid protoplast on the cell wall is called
turgor pressure (Fig.1.2). Since the cell wall is semi-elastic, it expands to a
certain extent. The cell wall prevents a plant cell from bursting, when placed
in water (hypertonic solution). An animal cell in a similar situation will burst.
The cell wall exerts an equal and opposite pressure to counter the turgor
pressure and it is called the wall pressure (WP). Thus, the TP and WP will be
equal but exerted in opposite directions Fig. 1.2).

TP = WP

Turgor pressure keeps the protoplast hydrated and appressed to the cell wall.
A fully (turgid) hydrated cell will have maximum hydrostatic pressure which will
be equal to its OP. On the other hand, TP = 0 in a flaccid cell. The value of TP
would, therefore, lie between zero and the value of OP. Under exceptional
18 conditions of plasmolysis, TP may even become negative.
Unit 1 Plant - Water Relation

Fig.1.2: Osmotic relations exhibited by a plant cell.

Turgor pressure helps in keeping the protoplasm in a hydrated state which is

important to maintain the 3-D structure of the organelles for their proper
functioning. In addition, turgor pressure helps in cell elongation, germination
and brings about a variety of plant movements including the seismonastic
movements as well as opening and closing of stomata.

1.3.6 Diffusion Pressure Deficit (DPD)

Each liquid possesses a definite diffusion pressure with which it can move.
Seismonastic
Pure water has maximum diffusion pressure. Addition of solute to it reduces its
movements are
diffusion pressure. So, the diffusion pressure of a solution is always less than paratonic movements
its pure solvent. This reduction in the diffusion pressure of water in a solution brought about by
or system over its pure state due to presence of solutes is called Diffusion external mechanical
Pressure Deficit (DPD). In other words, DPD is the difference between the stimuli like vibrations,
diffusion pressure of a solution and its pure solvent when present at the same strong winds, rain
drops, or contact with
temperature and pressure. The term DPD was coined by Meyer (1938). This
a foreign body.
term is preferred over the earlier term Suction Pressure (SP) proposed by
Renner (1915). DPD constitutes the power of absorption by the cell and
determines the direction of osmosis.

The earlier term SP was used to denote the amount of pressure by which
water can be sucked into the cell or likewise, expelled out of the cell. Another
way to define DPD will be that it is the amount of water needed by a cell to
make it fully turgid, as it is a measure of the water absorbing capacity of a cell.
A solution will absorb more water or solvent molecules to equalize its deficit of
diffusion pressure. So, water will always move from low DPD to high DPD.

Water Movement
Lower DPD Higher DPD

The value of DPD is always positive for a cell and is equal to osmotic pressure
of the system minus wall pressure.

DPD (SP) = OP – WP (=TP)

When a cell is placed in pure water or in a less concentrated solution

(hypotonic) than that of the cell sap, water enters the cell, resulting in an 19
Block 1 Water and Mineral Nutrition
increase in TP. Simultaneously the OP of the cell goes on decreasing as there
is a continuous inflow of water and the concentration of the cell sap goes
down. The value of TP goes on increasing and eventually becomes equal to
OP. At this stage, the cell becomes fully turgid

DPD = OP-TP

when OP = TP; then OP-TP = 0

DPD = 0.

Thus, a fully turgid cell does not have any capacity to suck water further.

On the other hand, when a cell is placed in a hypertonic solution, water from
the cell goes out by the process of exosmosis. As a result, the OP increases
and these in a corresponding decrease in TP. The cytoplasm contracts and
eventually the protoplast leaves the cell wall. This flaccid cell is said to be
plasmolyzed. At the time of complete plasmolysis, value of TP becomes zero.

So, at complete flaccidity, since TP = 0

and DPD (SP) = OP-TP

DPD = OP Such a cell will have maximum absorptive capacity.

If a cell was placed in some hypotonic solution other than water, then its

DPD (SP) = (OP-OP1) – TP (OP1, is osmotic pressure of the hypotonic

solution)

Sometimes the value of TP becomes negative when the cell wall of the
plasmolyzed cell pulls water in the opposite direction

Since TP is –ve

DPD = OP – (-TP)

DPD = OP + TP

Thus, the DPD of the plasmolyzed cell is greater than the value of OP. It is
clear from the above that the DPD of plant cell is not directly proportional to
osmotic concentration of the cell. It depends on OP as well as TP of the
osmotic system. So, it is DPD which is the basic determining force with which
a cell or an osmotic system will lose or absorb water, rather than the OP and
TP individually.

SAQ 2
In the following statements fill in the blank spaces with appropriate words:

a) Cells undergo plasmolysis when kept in ……………….solution.

b) Osmosis is diffusion of water through ………………. from a region of

high concentration of water to a low concentration of water.

c) In a hydrated cell, the hydrostatic pressure will be ………………, and

equal to its ………………
20
Unit 1 Plant - Water Relation
1.3.7 Chemical Potential
The chemical potential of a substance in a system is a measure of its free
energy. The free energy denotes the potential for performing useful work or
the energy available to do work, which is force x distance.

Here the system refers to the thermodynamic concept wherein studies are of
system rather than individuals or bodies. Thus, when we study the properties
of a solution in a 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 soil, 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 maximum,
and addition of any solute lowers the free energy or chemical potential of
water. This shows that the chemical potential is a relative quantity, which
represents the difference between the free energies (potential) of the
substance in each state to that substance in a standard state.

The unit of chemical potential is energy per mole of substance (J mol-1).

The chemical potential of water in a solution is related to is vapour pressure

and is given by the equation

μ −μ ° =  
°

∆ =  
°

Where:

µw = chemical potential of water in question (joules/mole)

µw° = chemical potential of pure water under STP

∆µ= change in free energy

R = gas constant

T = Absolute temperature in Kelvin

e = vapor pressure of water in question

Note that

Relative Humidity = ° × 100

If it is pure water, then  = 0 and ∆µ also becomes 0.
°
Partial Molal Volume
So, the chemical potential of water is set to zero. If chemical potential is less
 (J) of a solution is the
than that of pure water then  ° will be negative number, and therefore, ∆µ change in volume of a
will be less than zero, a negative number. So, the maximum value of chemical solution when one
potential is 0, and addition of the solute lowers it to negative values. Water will mole of a substance
spontaneously flow from regions of higher chemical potential to those is added to it.

possessing lower water potential. 21

Block 1 Water and Mineral Nutrition
1.3.8 Components of Water Potential
A related term Water potential (Stalyer and Taylor, 1960) has been used
Water potential is
historically by the plant physiologists, which is the difference between
expressed in pressure
units. Energy per unit chemical potential of water at any point in a system and that of pure water at
volume of water is STP (standard temperature and pressure). By conversion, the chemical
expressed in joules potential of water is referred to as water potential. It is denoted by the Greek
2
per cm . 106 dynes=1 letter Ψ (-pronounced as psi).
bar. The present unit
used for pressure is Ψ w is expressed in pressure units. It is the free energy per unit volume of
pascal (Pa). It is a water (Jcm-3). Ψw is equivalent to the chemical potential of water (∆µ) divided
pressure equal to the by the partial molal volume of water , i.e., the volume of 1 mol of water, which
force of one Newton is 18 × 10-6 m3 mol-1).
acting uniformly over
one square metre. If water potential of the source (the region supplying the water) in higher than
5 the water potential of the sink (the receiving region), then there is a
1 bar = 10 Pa
spontaneous transfer of water from the source to sink (Ψsource > Ψsink).
3
10 Pa= 1 KPa
(1 kilo pascal) By convention, the water potential of pure water at atmospheric pressure is
3 taken as zero. Therefore, water potentials other than that of pure water will
10 KPa = 1MPa
(mega pascal) generally be negative. Thus, lower potential means a more negative value,
and higher potential is less negative value.

Since the water potential of a system is related to the difference in the vapor
pressures of water in a solution and pure water state, it is taken as equivalent
to the diffusion pressure deficit (DPD).

As explained earlier, Ψw of pure water is 0, and so is the value of DPD.

Addition of solute (now a solution) will have a water potential as negative,
whereas the DPD would increase (greater than 0). You will read in the next
sections that the direction of movement of water is from lower DPD to higher
DPD. On the other hand, water moves from a region higher (less negative)
water potential to lower (more negative) water potential.

Water potential in plants is affected by three major factors: solute

concentration, hydrostatic pressure, and matric forces. In addition, gravity also
plays some role. Water potential (Ψ/Ψw) is equal to the algebric sum of the
potentials – Solute potential (Ψs), pressure potential (Ψp), matric potential
(Ψm) and gravitational component (Ψg – when present).

Ψw = Ψs +Ψp + Ψm + Ψg

Let us consider each of these components in details:

Effect of Solute
Let us take pure water in two chambers A and B separated by a semi-
permeable membrane (Fig. 1.3a). The water potential of both the chamber is
zero. Addition of solute in chamber A (Fig. 1.3b) would reduce the free energy
of water and the water potential will fall below zero. Consequently, water will
move from B to A (Fig. 1.3c). The effect of dissolved solutes on water potential
(Ψw) is called osmotic potential (Ψπ). It can be estimated numerically if we
know the osmotic pressure of the solution. The two are related as

π = –Ψπ

22 For example, if π of solution is 5 bars then Ψπ would be –5 bars.

Unit 1 Plant - Water Relation

Osmolality = moles
of total dissolved
solutes per litre of
-1
water – (mol L ). 6.02
23
× 10 particles of the
non-ionic solute will
be present in one litre
of solution with -22.7
bars at O°C = 1 mole
of non-electrolyte and
-48.3 bars for the
electrolyte sodium
chloride.

Fig.1.3: Experiment to show the effect of solute and pressure on water potential.

In other words, osmotic potential or solute potential (Ψs) represents impact of

dissolved solutes on its water potential i.e.it records the decrease in the water
potential or free energy of water due to the presence of solute particles (ionic
or non-ionic). Under ideal conditions, the Ψs is also equivalent to the
maximum osmotic pressure which can be developed under ideal conditions
but written with a negative sign as the dissolved solutes reduce the Ψw of a
solution with reference. Dilute non-ionic solutions, e.g., sucrose and glucose
will have an osmotic potential (solute potential).

Ψs = –RTCs (see osmotic pressure Section 1.3.5 for values of R and T)

Osmotic potential of 1.0 molal solution of sorbitol at 20°C will be [Ψs = –ci RT]
: –(1.0mol kg-1) (1.0) (0.00831 kg MPa mol-1 k-1/0.080205 kg atm mol-1 k-1) (20
+ 273 = 293°K)= 2.43 MPa]

The solute potential of the electrolyte will decrease by the degree of its
dissociation over that of a non-electrolyte. To be expressed as Ψs (electrolyte)
= iRT. By this analogy, 0.33 M calcium chloride solution will have the same
solute potential as 1.0 M sucrose.

Effect of Pressure

Let us now see the effect of pressure on water potential. As Fig. 1.3d
illustrates, when pressure is applied the flow of water begins from chamber A 23
Block 1 Water and Mineral Nutrition
to chamber B 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 Ψp. The level of water in B rises due to
increase in water potential of A (Fig. 1.3 e). Pressure potential is usually
positive (is equal to turgor pressure). The values of turgor pressure are
particularly high in leaf cells of crop plants during warm afternoons and in
guard cells during stomatal opening.

In sharp contrast, xylem and in the walls between cells, a negative hydrostatic
pressure or tension develops, leading to the value of Ψp becoming negative.

Now see the Fig. 1.3 where two chambers are present. What would happen if
pressure were applied on the chamber containing solutes (Fig.1.3 d)? Now the
Ψw 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 (Fig.1.3 e). An equilibrium condition will
reach when pressure potential Ψp will be equal but opposite in magnitude to
osmotic potential Ψπ and there will no net flow of water in the two chambers
(Fig.1.3 f). This can be represented by the following equation:

Ψw(B) = (Ψπ + Ψp)B …(1.1)

Let us suppose if Ψp is equal and opposite to Ψπ. Then
ΨWA = (ΨπA + ΨπB)
=0
We are going to discuss another important factor, i.e. the “Matric pressure”.

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 (Ψm). Its value is always negative.

Matric potential can be viewed as a decrease in water potential on account of

immobilization of water molecules due to their adsorption over the colloidal
particles in the cytoplasm and cell wall. The value of Ψm may be appreciable in
young cells, dry seeds, fruits, and some extreme xerophytes. The mature cells
of mesophytes, however, show an extremely low Ψm value of –0.1 bars, and
thus, can be ignored.

Likewise, in a well-watered soil Ψm is not very significant, however, when the

soil is nearly drying Ψm determines water potential of the soil.

It is very natural that the water column will be forced to move downward due to
the gravitational force. Thus height, density of water and the acceleration due
to gravity will play a role to decide the equal and opposite force to counter this
gravitational potential– Ψg. It has been calculated that the value of Ψg is
quite insignificant (accounting for nearly 0.1 MPa in water potential) and is,
24 therefore, generally not considered for water transport at the cell level.
Unit 1 Plant - Water Relation
So, we find that the Ψm is composed of the following main component forces:

Ψw= Ψπ+Ψp + Ψm+ Ψ … (1.2)

Ψ= any other forces that may influence Ψw. Thus, water potential is equal to
Ψw= Ψs= Ψp

Resistances to Water Movement and Water Flux

We have explained earlier that if water potential drops from source to sink st
Fick’s 1 Law :
(Ψsource>Ψsink) then there will be spontaneous flow of water. But we do not Diffusive flux =
know the rate of this transfer i.e., flux. Let us learn how to calculate flux. diffusion coefficient;
concentration
Flux describes any effect that appears to pass or travel (whether it moves or gradient
not) through a surface or substance. With reference to the transport
J = Ds∆/∆x
phenomena, flux is considered as a vector quantity (having both magnitude
J = Flux rate
and direction) which describes the magnitude and direction of the flow of a 2
(moles/m )
substance. The diffusive flux can be related to the concentration gradient
using Fick's first law, which postulates that the flux moves from regions of ∆Cs/∆x is
concentration
high concentration to regions of low concentration. The magnitude of flux is
gradient
proportional to the concentration gradient. In other words, the solute moves -3
(moles m /m)
from a region of high concentration to a region of low concentration across a
Ds = diffusion
concentration gradient. 2 -1
coefficient (m s )
The flux for water flow is denoted by Jw which 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 = Lp ∆Ψw … (1.3)

Lp = permeability coefficient of limiting membranes

∆Ψw = difference in water potential at two points

From equation (1.1) we know that

Ψw = (Ψπ + Ψp)

∆Ψw = (∆Ψπ + ∆Ψp)

substituting the value of ∆Ψw in above equation (1.3)

Jw = Lp(∆Ψπ + ∆Ψp) …(1.4)

Thus, the inward or outward rate of flow or water from cell to cell and tissues
can be calculated from the above expression.

The water flux is directly proportional to ∆Ψ and inversely proportional to

hydraulic resistance (R). In other words, higher the ∆Ψw ,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
exert more resistance (because of low permeability) than the cell walls. 25
Block 1 Water and Mineral Nutrition
Therefore, water can flow relatively easily through cell walls. Water will not
Hydrostatic
experience the resistance of plasma membrane when it moves from cell to cell
pressure is the
pressure exerted by via plasmodesmata. The xylem conduits which are not obstructed by cell
or on a liquid above membranes have least resistance and the rate of flow of water is very high.
or below atmospheric The ratio of resistance in xylem, cell walls and cell membranes is in the order
pressure. of 0.3:1: 50. This 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 ∆Ψw in soil is determined by the matric pressure.

∆Ψw(soil) = – ∆Ψm(soil)

SAQ 3
a) List the three factors that determine the value of Ψw in plant.
b) The water potential in a cup (Ψc) containing salt solution will be
Ψc>Ψw

Ψc<Ψw

Ψc= Ψw

c) What will be the water potential of a plasmolyzed 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 Ψw of a cell will be……………….
ii) The net flow of water movement in system will stop when Ψp will
be equal and ………………… to Ψπ.
iii) Greater matric suction will ……………… the water potential in a
system.

1.3.9 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 organization. We have already discussed
the long-distance transport of water.

Movement 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 up to 90% of the cell and contains cell sap which is a
dilute solution of salt and other small molecules. Due to osmotic potential the
26 cell sap has a lower water potential than pure water. When such a cell is
Unit 1 Plant - Water Relation
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. 1.4).
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:

Ψw (outside the cell) = (Ψπ + Ψp) inside the cell

where, Ψw is total water potential of the system, Ψπ, is the osmotic or solute
potential and Ψp, is pressure potential due to cell wall pressure or turgor
pressure. At full turgor pressure the sum of Ψp, and Ψπ, is zero. Hence Ψw is
zero.

The driving force F that causes water to move can be represented by the
following equation:

F = gradient (Ψc –Ψe)

Where Ψc is the total water potential of the cell including that of the cell sap
and Ψe 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
Ψc. However, as the water will move into the cell Ψc will be regulated by Ψπ
and Ψp. 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.

Fig. 1.4: A single cell surrounded by water (diagrammatic).

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
apoplast is determined by matric forces exerted in cell wall.

It must be now clear to you that the osmotic status of the cell changes in
response to gain or loss of water. The Fig.1.5 below (called Hofler diagram)
illustrates the relationship between Ψw, Ψs, and Ψp (along with DPD, OP and
WP) in response to gain or loss of water. 27
Block 1 Water and Mineral Nutrition

Fig.1.5: Hofler diagram depicting the relationship between Ψw , Ψs , and Ψp

(along with DPD, OP and WP)
Placed in a hypotonic solution, endosmosis causes the expansion of the
central vacuole, thereby swelling the protoplast and a rise in turgor pressure or
pressure potential. At full turgidity turgor pressure (TP) rises to a level which is
equal to osmotic pressure(OP). CurrentlyΨw or diffusion pressure deficit (DPD)
becomes zero, and there is no further entry of water into the cell

DPD = OP-TP Ψw = (-)Ψs–(+)Ψp

Ψw = Ψs + Ψp
At full turgidity TP = OP [Ψs= Ψp]
DPD = 0 Ψw = 0 (A)

Alternatively, when a turgid cell is immersed in a hypertonic solution, loss of

water results. Due to water loss the solute potential of the cell sap becomes
slightly more negative (slight increase in OP). The pressure potential or turgor
pressure decreases and at full flaccidity, becomes zero. At this point the water
potential shows maximal decline to equal to osmotic potential. (B)

In a flaccid cell

DPD = OP-TP Ψw = (-)Ψ s – (+)Ψ p

Ψw = Ψs + Ψp
Since TP = 0 Since Ψp = 0
DPD = OP Ψw = Ψs

Water Relations of a Tissue

In higher plants no cell exists in isolation separate from others. Even a root
hair which is projected outside into the surrounding medium is attached with
other cells on all there maining 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
28 them. This can be shown as follows:
Unit 1 Plant - Water Relation
The total water potential of cell A (ΨwA) will be equal to the sum of its osmotic
potential and pressure potential (ΨπA+ ΨpA). Let us say that for the cells A and
B,these values are

ΨwA = (ΨπA+ ΨpA)

ΨwB = (ΨπB+ ΨpB)

In a cell the matric forces are much less. Here the Ψw is lowered because of
Ψπ. If the Ψw in the vacuole is lower than Ψw in apoplast, then water flows
inwards.

Now, the driving force (F) will be the difference in the water potential of two
cells.

F = (ΨwA– ΨwB)

If ΨwA is equal toΨwB the gradient will be zero and so will be the driving force
(F). Therefore, no net exchange of water will take place between the cells A
and B. However, you must realize that the same value of ΨwA and ΨwB does
not necessarily mean that the two cells should have the same osmotic
potential and the same turgor pressure or wall pressure.

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 1 to 20, they will attain an
equilibrium amongst themselves, depending 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.

To illustrate this point by an example, if two adjacent cells A and B have the
following values. Cell A has an osmotic potential (Ψs) of–9 bars and its
pressure potential (Ψp) is equivalent to 6 bars, while the cell B has its
corresponding values of –12 bars and 4 bars respectively, the flow of water
will be :

Ψw= Ψs+ Ψp
A B
Ψw= –9 +6 Ψw= –12 + 4

= –3 bars = –8 bars

Since water moves from less negative (higher) Ψw to more negative (lower)
Ψw, movement of water will be from cell A to cell B.

Water Relations of a Whole Plant

Let us now consider the water relations of a plant considering leaves, stem,
and roots, that provide a continuum for water in soil-plant-atmosphere system.
The total water potential in the atmosphere could be very low, depending upon
the temperature and humidity. Table 1.1 shows an approximate magnitude of
water potential in the soil-plant-atmosphere-system. 29
Block 1 Water and Mineral Nutrition
Table.1.1: Water Potential.

Component Water Potential (Bars)

Soil -0.1 to -20.0

Leaf -5.0 to -50.0
Atmosphere -100 to -2000

It is clear from the data in Table 1.1 that the difference in total water potential
in 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.

1.4 SUMMARY
• Water is the key molecule for the maintenance of life on the earth. Plant
has about 85 to 90% 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 can 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 cohesive 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 (Ψw). 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 algebric sum of component
forces: osmotic potential (due to solute), pressure potential (due to
hydrostatic pressure) matric or 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
30 along the cell walls.
Unit 1 Plant - Water Relation

1.5 TERMINAL QUESTIONS

1. Differentiate between:

a) Diffusion and Osmosis

b) Osmotic Potential and matric potential

c) Apoplastic and symplastic pathway

d) Imbibition and plasmolysis

2. Write brief notes on:

a) Aquaporins

b) Units of water potential

c) The concept of water potential and its components

d) Unique properties of water

3. What is water potential? Discuss its components and derive a

relationship between them with the help of Hofler diagram.

4. Explain how the osmotic pressure vary in electrolytes and non-

electrolytes.

1.6 ANSWERS
Self-Assessment Questions
1. a) Heat of vaporization

b) Cohesion

c) 10 picoseconds

d) Dipoles

e) Germination

f) 0.239 calorie

2. a) hypertonic

b) semi-permeable membrane;

c) maximum, Osmotic potential.

3. a) Osmotic potential (Ψπ), pressure potential (Ψp) and matric

potential (Ψm).

b) Ψp= Ψw the Ψc of salt solution in a cup will be zero because there

are not membranes present. Therefore, its water potential is equal
to pure water.

c) -7.9 MPa.

We know that π = Ψπ. Therefore, the osmotic potential will be -7.9

MPa. Since Ψp= 0, therefore, Ψw = Ψπ. 31
Block 1 Water and Mineral Nutrition
d) i) zero

ii) opposite

iii) lower

Terminal Questions
1. a) Refer to Section 1.3.

b) Refer to Subsection 1.3.2.

c) Refer to Section 1.4.

d) Refer to Subsection 1.3.2 and 1.3.8.

2. a) Refer to Subsection 1.3.4.

b) Refer to Subsection 1.3.8.

c) Refer to Subsection 1.3.8.

d) Refer to Section 1.2.

3. Refer to Subsection 1.3.8 and 1.3.9.

4. Refer to Subsection 1.3.5.

32