Transport in Plants

Introduction:
 Plants need to move molecules over very long distances, much more than animals do;
they also do not have a circulatory system in place.
 Water taken up by the roots has to reach all parts of the plant, up to the very tip of the
growing stem.
 The photosynthates or food synthesised by the leaves have also to be moved to all parts
including the root tips embedded deep inside the soil.
 Movement across short distances, say within the cell, across the membranes and from cell
to cell within the tissue has also to take place.
 In a flowering plant the substances that would need to be transported are water, mineral
nutrients, organic nutrients and plant growth regulators.
 Over small distances substances move by diffusion and by cytoplasmic streaming
supplemented by active transport.
 Transport over longer distances proceeds through the vascular system (the xylem and the
phloem) and is called translocation.
 An important aspect that needs to be considered is the direction of transport.
 In rooted plants, transport in xylem (of water and minerals) is essentially unidirectional,
from roots to the stems.
 Organic and mineral nutrients however, undergo multidirectional transport.
 Organic compounds synthesised in the photosynthetic leaves are exported to all other
parts of the plant including storage organs.
 The mineral nutrients are taken up by the roots and transported upwards into the stem,
leaves and the growing regions.
 When any plant part undergoes senescence, nutrients may be withdrawn from such
regions and moved to the growing parts.
 Hormones or plant growth regulators and other chemical stimuli are also transported,
though in very small amounts, sometimes in a strictly polarised or unidirectional manner
from where they are synthesised to other parts.
 Hence, in a flowering plant there is a complex traffic of compounds (but probably very
orderly) moving in different directions, each organ receiving some substances and giving
out some others.

Means of Transport:
 The transport of material into and out of the cells is carried out by a number of methods.
These are diffusion, facilitated diffusion and active transport.
Diffusion :
 Diffusion occurs from region of higher concentration to region of lower concentration
across the permeable membrane. It is passive and slow process. No energy expenditure
takes place.
 Factors affecting diffusion:
 Concentration gradient,
 Permeability of membrane; separating the substances
 Temperature and Pressure

Facilitated Diffusion:
 While non-polar substances diffuse through the membrane, the polar substances move
with help of special proteins. This process is called facilitated diffusion.
  The diffusion of hydrophilic substances along the concentration gradient through fixed
membrane transport protein without involving energy expenditure is called facilitated
diffusion. For this the membrane possesses aquarporins and ion channels. No energy is
utilized in this process.
 Carrier proteins form porin channels in the outer membranes of plastids, mitochondria
and some bacteria. The porin channels allow the molecules up to the size of small
proteins to pass through molecule bound to the transport protein. The transport protein
then rotates and releases the molecule inside the cell. For example; water channels are
made up of eight different types of aquaporins.

Symports and Antiports:

 Some carrier proteins allow diffusion only if two types of molecules move together.
 When both the molecules cross the membrane in the same direction, it is called symport.
 When the molecules move in opposite directions, it is called antiport.
 When a molecule moves across a membrane independent of other molecules, the
movement is called uniport.
Active Transport:
 Energy pumps are used against a concentration gradient; in case of active transport.
 Active transport is carried out by membrane proteins. Pumps are proteins which use
energy to carry substances across the cell membrane.
 The rate of transport reaches the maximum when all the protein transporters are being
used or are saturated.

Plant Water Relations:

 Water is essential for all physiological (metabolic) activities of all living organisms
including plants without which they cannot survive as most of the metabolites are are
dissolved in it.
 Protoplasm is made up of water in which various metabolites are dissolved & suspended.
 Distribution of water within a plant varies – wherein soft parts mostly contain water
while woody parts have relatively very little water.
 Seeds also contain small amount of water even though they appear dry.
 Land plants take up huge amount of water daily but most of it is lost to the air through
evaporation from the leaves, i.e., transpiration.
 Water is often the limiting factor for plant growth and productivity in both agricultural
and natural environments due to high demand of water by them.
Water Potential:
 Water molecules possess kinetic energy.
 A system with higher concentration of water has a higher kinetic energy or water
potential.
 Water potential is the chemical potential of water.
 Water potential of a system is defined as the difference between the free energy of water
in the system & the free energy of pure water.
 Pure water has the highest water potential.
 Solutions have lower water potential than pure water.
 If two systems containing water are in contact, random movement of water molecules
will result in net movement of water molecules from the system with higher energy to the
one with lower energy.
 Movement happens from higher water potential to lower water potential.
 This process of movement of substances down a gradient of free energy is called
diffusion.
 Water potential is denoted by Ψw (Psi) and measured in Pascals (Pa).
 Water potential of pure water at standard temperature which is not under any pressure is
taken to be zero (by convention).
Solute Potential:
o All solutions have a lower water potential because of the dissolved solutes.
o The magnitude of lowering of water potential is called solute potential (Ψs).
o Solute potential is always negative.
o For a solution at atmospheric pressure:
Water potential = Solute potential
Pressure Potential:
o If a pressure greater than atmospheric pressure is applied to pure water or a solution, its
water potential increases.
o This happens because of pressure potential which develops due to increased pressure.
o Pressure potential is usually positive. Pressure potential is expressed as Ψp
The water potential of a cell is affected by solute potential (Ψs) and pressure potential (Ψp).
The relationship between them is as follows: Ψw = Ψs + Ψp

Osmosis:
o Diffusion of water across a semipermeable membrane is called osmosis.
o The net direction and rate of osmosis depends on pressure gradient and concentration
gradient.
o Water moves from higher concentration to its lower concentration until equilibrium is
achieved.
o The two chambers across the semi-permeable membrane have the same water potential at
equilibrium.
o When a solution of sucrose in water taken in a funnel is separated from pure water in a
beaker through a semi-permeable membrane, water will move into the funnel, resulting in
rise in the level of the solution in the funnel. This will continue till the equilibrium is
reached.
o External pressure can be applied from the upper part of the funnel such that no water
diffuses into the funnel through the membrane.
o This pressure required to prevent water from diffusing is in fact, the osmotic pressure and
this is the function of the solute concentration; more the solute concentration, greater will
be the pressure required to prevent water from diffusing in.
o Numerically osmotic pressure is equivalent to the osmotic potential, but the sign is
opposite.
o Osmotic pressure is the positive pressure applied, while osmotic potential is negative.
Kinds of Osmosis:
There are two types of osmosis viz. endosmosis & exosmosis
 Endosmosis – is the entry of water into the cell through a semipermeable membrane,
wherein the cells become turgid.
 Exosmosis – is the exit of water from the cell to the external environment resulting in the
cell becoming flaccid’
Significance of Osmosis:
1. Root hairs absorb water by endosmosis. The cellular water enters the central vacuole by
osmosis, through vacuolar membrane or tonoplast.
2. Intercellular movement of water is through osmosis.
3. Cells become turgid due to the entry of water through endosmosis. Turgidity of the ell is
necessary for physiological activities.
4. Opening & closing of stomata is due to endosmosis & exosmosis which is necessary for
gaseous exchange & transpiration.

Plasmolysis:
o Depending on the nature of surrounding solution, the cells behave differently to water
movement.
o If the external solution balances the osmotic pressure of the cytoplasm, it is said to be
isotonic.
o If the external solution is more dilute than the cytoplasm, it is hypotonic and if the
external solution is more concentrated, it is hypertonic.
o When a plant cell is placed in hypertonic solution, the plant cell loses water and hence it
loses the turgor pressure.
o This makes the cell flaccid.
o The plant cells wilt in this condition.
o Further water loss results in plasmolysis.
o At this point, the pressure decreases to an extent where the protoplasm of the cell peels
away from the cell wall.
o This leaves gaps between the cell wall and the membrane.
o This can also lead to cytorrhysis or complete collapse of the cell wall.
o When the cell (or tissue) is placed in an isotonic solution, there is no net flow of water
towards the inside or outside.
o If the external solution balances the osmotic pressure of the cytoplasm it is said to be
isotonic.
o Plasmolysis can be reversed by putting the cell in hypotonic solution.
o When the cells are placed in a hypotonic solution (higher water potential or dilute
solution as compared to the cytoplasm), water diffuses into the cell causing the cytoplasm
to build up a pressure against the wall, that is called turgor pressure.
o This leads to the cell getting swollen.
o The pressure exerted by the protoplasts due to entry of water against the rigid walls is
called pressure potential Ψp.
o But the rigidity of the cell wall prevents the cell from bursting.
o Plasmolysis rarely happens in nature and can only occur in extreme conditions.

Imbibition:
o Imbibition is the phenomenon of adsorption of water or any other liquid by the solid
particles of a substance without forming a solution.
o It is a special type of diffusion when water is absorbed by solids – colloids – causing
them to enormously increase in volume.
 o Imbibition is also diffusion since water movement is along a concentration gradient; the
seeds and other such materials have almost no water hence they absorb water easily.
Some examples of Imbibition :
 If a dry piece of wood is placed in water, it swells and increases in its volume.
 If dry gum or pieces of agar-agar are placed in water, they swell and their volume
increases.
 When seeds are placed in water they swell up.

Long Distance Transport in Plants:

 Mass flow: Mass flow is the movement of substances (water, minerals and food) in bulk
from one point to another as a result of pressure differences between two points.
 It is a characteristic of mass flow that substances, whether in solution or in suspension,
are swept along at the same pace, as in a flowing river.
 This is unlike diffusion where different substances move independently depending on
their concentration gradients.
 Bulk flow can be achieved either through a positive hydrostatic pressure gradient (e.g., a
garden hose) or a negative hydrostatic pressure gradient (e.g., suction through a straw).
 Translocation: the bulk movement of substance through the conducting or vascular
tissue is called translocation.
Absorption of Water:
 Absorption of water from the soil happens through root hairs.
 Root hairs are extensions of the root epidblema and have thin walls. Water enters the root
hairs because of osmosis.
 Presence of numerous root hairs increases the surface area and hence enhances the
absorption of water.
 Water is absorbed by root hairs, then water moves upto xylem by two pathways −
apoplast and symplast pathway.
Apoplast pathway :
 Movement of water takes place exclusively through the intercellular spaces and the walls
of the cells.
 Movement through the apoplast does not involve crossing the cell membrane.
 Movement depends on the gradient.
 The apoplast does not provide any barrier to water movement.
 Water movement is through mass flow.
Symplast pathway :
 System of interconnected protoplasts.
 Neighboring cells are connected through cytoplasmic strands that extend through
plasmodesmata.
 Water enters into the cytoplasm by crossing the plasma membrane.
 Intercellular movement is through the plasmodesmata.
  Endodermis is impervious to water because of a band of suberised matrix called
casparian strip.
 Water molecules are unable to penetrate the layer, so they are directed to wall regions
that are not suberised, into the cells proper through the membranes.
 The water then moves through the symplast and again crosses a membrane to reach the
cells of the xylem.
 The movement of water through the root layers is ultimately symplastic in the
endodermis.
 This is the only way water and other solutes can enter the vascular cylinder.
 Once inside the xylem, water is again free to move between cells as well as through them.
 In young roots, water enters directly into the xylem vessels and/or tracheids. These are
non-living conduits and so are parts of the apoplast.

Root Pressure:
 When various ions from the soil are actively transported into the vascular tissues of the
roots, water also follows.
 This increases the pressure inside the xylem.
 This positive pressure is called root pressure.
 The root pressure can push water up to small heights in the stem.
Guttation:
 In some plants, under the conditions of low evaporation, water comes out from the tips of
leaves.
 Such loss of water in its liquid phase is called guttation.
 Guttation takes place in smaller plants only.
Limitations of Root Pressure:
 Root pressure can only provide a modest push.
 It does not play a major role in water movement in tall plants.
 Root pressure contributes towards reestablishment of continuous chains of water
molecules in the xylem; which often break under enormous tensions created by
transpiration pull.
Transpiration Pull:
 The transport of water to the tops of trees occurs through xylem vessels.
 The forces of adhesion and cohesion maintain thin and unbroken columns of water in
the capillaries of xylem vessels through which it travels upward.
 Water is mainly pulled by transpiration from leaves. (Cohesion-tension-transpiration
pull Model)
 Transpiration creates a suction force inside the xylem.
 This suction force is called transpiration pull.
  This is powerful enough to pull the water column from beneath.
 Adhesion, cohesion and surface tension are the important physical properties of water
which further help in the upward movement of water through xylem.
 Cohesion: Mutual attraction between water molecules is called cohesion.
 Adhesion: Attraction of water molecules to polar surfaces is called adhesion.
 Surface Tension: Any liquid has a tendency to occupy the least possible surface area.
This property is called surface tension.
 The above mentioned properties impart high tensile strength to water. The high tensile
strength imparts an ability to resist a pulling force and high capillarity i.e., the ability to
rise in thin tubes. The thin tubes of xylem work like capillary tubes.
 In plants capillarity is aided by the small diameter of the tracheary elements – the
tracheids and vessel elements.

Transpiration:
The loss of water through stomata of leaves and other aerial parts of plants in form of water
vapour is known as transpiration.
o Transpiration mainly occurs through stomata.
o Stomata are usually open during daytime and remain closed during the night.
Opening and Closing of Stomata:
o A change in the turgidity of guard cells results in closing or opening of stomata.
o The inner wall of the guard cell, towards the stomatal aperture is thick and elastic.
o An increase in turgidity results in the thin outer walls to bulge out.
o This forces the inner wall into a crescent shape and results in opening of stoma.
o The orientation of the microfibrils in the cell walls of the guard cells also helps in
opening of stomata.
o These microbifibrils are radially oriented and thus make it easy for the stoma to open up.
o A loss in turgidity of the guard cells, leads to resumption of shape of the elastic inner wall
of the guard cell and the stoma closes.
o The process of photosynthesis requires water.
o The system of xylem vessels from the root to the leaf vein can supply the needed water.
o As water evaporates through the stomata, since the thin film of water over the cells is
continuous, it results in pulling of water, molecule by molecule, into the leaf from the
xylem.
o Also, because of lower concentration of water vapour in the atmosphere as compared to
the substomatal cavity and intercellular spaces, water diffuses into the surrounding air.
This creates a ‘pull’.
o Measurements reveal that the forces generated by transpiration can create pressures
sufficient to lift a xylem sized column of water over 130 metres high.
Factors Affecting Transpiration:
o Temperature, light, humidity, wind speed are the external factors affecting transpiration.
o Plant factors affecting transpiration are number and distribution of stomata, number of
stomatal aperture with guard cells open, water status of the plant, canopy structure, etc.

Role of transpiration :
 Creates transpiration pull for absorption and transport of plants.
 Supplies water for photosynthesis.
 Transports minerals from the soil to all parts of the plants.
 Cools leaf surfaces, sometimes 10 to 15 degrees, by evaporative cooling.
 Maintains the shape and structure of the plants be keeping cells turgid.

Uptake & Transport of Minerals:

Minerals cannot be passively absorbed by roots. There are two main reasons for this:
 Minerals are present as charged particles in soil. They cannot move across cell
membranes.
 Concentration of minerals in the soil is usually lower than the concentration of minerals
in the root.
Hence, minerals need to be actively absorbed by the epidermal cells, which needs energy in the
form of ATP. Specific proteins in the membranes of root hairs actively pump ions from the soil
to the epidermal cells. Transport proteins of endodermal cells are control points, where a plant
adjusts the quantity and types of solutes that reach the xylem. The root endodermis because of
the layer of suberin has the ability to actively transport ions in one direction only.

Translocation of Mineral Ions:

 Minerals ions reach xylem through active or passive uptake, or a combination of both.
 Their further movement through the xylem is along with the transpiration stream.
 The growing regions of the plant are the main sinks for mineral elements.
 Mineral ions are frequently mobilized; especially from older, senescing parts.
 Phosphorus, sulphur, nitrogen and potassium are the most readily mobilized elements.
 However, elements which are structural components are not mobilized, e.g. calcium.
Some elements are also transported in organic forms in plants.

Phloem Transport:
 Food is transported through phloem; from source to sink.
 Leaf usually plays the role of source and storage organs are the sinks.
 But there can be role reversal when new leaves emerge during early spring.
 Thus, movement of substances through phloem is bi-directional.
 The phloem sap is mainly composed of water and sucrose, but other sugars, hormones
and amino acids may also be present.
The Pressure Flow Or Mass Flow Hypothesis:
 When glucose is prepared at the source, it is converted to sucrose.
 The sucrose moves into the companion cells and then into the living phloem sieve tube
cells; through active transport. This process of loading at the source produces a
hypertonic condition in the phloem.
 Water; from the adjacent xylem; moves into the phloem, by osmosis. This results in an
increase of osmotic pressure. It forces the phloem sap to areas of lower pressure, i.e.
towards the sink. The osmotic pressure must be reduced at the sink.
 Active transport moves the sucrose out of the phloem sap into the cells in the sink. Once
the sugar is removed, the osmotic pressure decreases and water moves out of the phloem.