Transport in plants,higher secondary important topics for various examination like NEET,AFMC,AIMS,board examinations…we provide pdf,notes,questions,videos of this chapter
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. From the storage organs
they are later re-exported. 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 When any plant part undergoes senescence, nutrients may be withdrawn from such regions and moved to the growing parts.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 signals 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.
11.1 MEANS OF TRANSPORT
11.1.1 Diffusion
Movement by diffusion is passive, and may be from one part of the cell to
the other, or from cell to cell, or over short distances, say, from the intercellular spaces of the leaf to the outside. No energy expenditure takes place.
In diffusion, molecules move in a random fashion, the net result being substances moving from regions of higher concentration to regions of lower
concentration. Diffusion is a slow process and is not dependent on a ‘living
system’. Diffusion is obvious in gases and liquids, but diffusion in solids
rather than of solids is more likely. Diffusion is very important to plants
since it is the only means for gaseous movement within the plant body.
Diffusion rates are affected by the gradient of concentration, the permeability of the membrane separating them, temperature and pressure.
Related:Animal kingdom
11.1.2 Facilitated Diffusion
As pointed out earlier, a gradient must already be present for diffusion to
occur. The diffusion rate depends on the size of the substances; obviously
smaller substances diffuse faster. The diffusion of any substance across a membrane also depends on its solubility in lipids, the major constituent of
the membrane. Substances soluble in lipids diffuse through the membrane
faster. Substances that have a hydrophilic moiety, find it difficult to pass
through the membrane; their movement has to be facilitated. Membrane
proteins provide sites at which such molecules cross the membrane. They
do not set up a concentration gradient: a concentration gradient must
already be present for molecules to diffuse even if facilitated by the proteins.This process is called facilitated diffusion.
In facilitated diffusion special proteins help move substances across membranes without expenditure of ATP energy. Facilitated diffusion
cannot cause net transport of molecules from a low to a high concentration
– this would require input of energy. Transport rate reaches a maximum
when all of the protein transporters are being used (saturation). Facilitated
diffusion is very specific: it allows cell to select substances for uptake. It is
sensitive to inhibitors which react with protein side chains.
The proteins form channels in the membrane for molecules to pass through. Some channels are always open; others can be controlled. Some are large, allowing a variety of molecules to cross. The porins are proteins that form large pores in the outer membranes of the plastids, mitochondria and some bacteria allowing molecules up to the size of small proteins to pass through. shows an extracellular A molecule bound to the transport protein; the transport protein then rotates and releases the molecule inside the cell, e.g., water channels – made up of eight different types of aquaporins.
11.1.2.1 Passive symports and antiports
Some carrier or transport proteins allow diffusion only if two types of molecules move together. In a symport, both molecules cross the membrane in the same Membrane direction; in an antiport, they move in opposite directions . When a molecule moves across a membrane independent of other molecules, the process is called uniport.
11.1.3 Active Transport
Active transport uses energy to transport and pump molecules against a concentration gradient. Active transport is carried out by specific membrane-proteins. Hence different proteins in the membrane play a
major role in both active as well as passive transport. Pumps are proteins
that use energy to carry substances across the cell membrane. These
pumps can transport substances from a low concentration to a high concentration (‘uphill’ transport). Transport rate reaches a maximum
when all the protein transporters are being used or are saturated. Like
enzymes the carrier protein is very specific in what it carries across the membrane. These proteins are sensitive to inhibitors that react with protein
side chains.
11.1.4 Comparison of Different Transport Processes
Proteins in the membrane are responsible for facilitated diffusion and
active transport and hence show common characterstics of being highly
selective; they are liable to saturate, respond to inhibitors and are under
hormonal regulation. But diffusion whether facilitated or not – take place
only along a gradient and do not use energy.1.2 PLANT-WATER RELATIONS
Water is essential for all physiological activities of the plant and plays a
very important role in all living organisms. It provides the medium in
which most substances are dissolved. The protoplasm of the cells is
nothing but water in which different molecules are dissolved and (several
particles) suspended. A watermelon has over 92 per cent water; most herbaceous plants have only about 10 to 15 per cent of its fresh weight
as dry matter. Of course, distribution of water within a plant varies –
woody parts have relatively very little water, while soft parts mostly contain water. A seed may appear dry but it still has water – otherwise it would not be alive and respiring! Terrestrial plants take up huge amount water daily but most of it is lost to the air through evaporation from the leaves, i.e., transpiration. A mature corn plant absorbs almost three litres of water in a day, while a mustard plant absorbs water equal to its own weight in about 5 hours. Because of this high demand for water, it is not surprising that water is often the limiting factor for plant growth and productivity in both agricultural and natural environments.
11.2.1 Water Potential
To comprehend plant-water relations, an understanding of certain
standard terms is necessary. Water potential (Ψw) is a concept
fundamental to understanding water movement. Solute potential
(Ψs) and pressure potential ( Ψp) are the two main components that
determine water potential. Water molecules possess kinetic energy. In liquid and gaseous form they are in random motion that is both rapid and constant. The greater the concentration of water in a system, the greater is its kinetic energy or ‘water potential’. Hence, it is obvious that pure water will have the greatest water potential. 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. Thus water will move from the system containing water at higher water potential to the one having low water potential. This process of movement of substances down a gradient of free energy is called diffusion. Water potential is denoted by the Greek symbol Psi or Ψ and is expressed in pressure units such as pascals (Pa). By convention, the water potential of pure water at standard temperatures, which is not under any pressure,is taken to be zero. If some solute is dissolved in pure water, the solution has fewer free water molecules and the concentration (free energy) of water decreases, reducing its water potential. Hence, all solutions have a lower water potential than pure water; the magnitude of this lowering due to dissolution of a solute is called solute potential or Ψs. Ψs is always negative. The more the solute molecules, the lower (more negative) is the Ψs . For a solution at atmospheric pressure (water potential) Ψw = (solute potential) Ψs. If a pressure greater than atmospheric pressure is applied to pure water or a solution, its water potential increases. It is equivalent to pumping water from one place to another. Pressure can build up in a plant system when water enters a plant cell due to diffusion causing a pressure built up against the cell wall, it makes the cell turgid ,this increases the pressure potential. Pressure potential is usually
positive, though in plants negative potential or tension in the water column
in the xylem plays a major role in water transport up a stem. Pressure
potential is denoted as Ψp. Water potential of a cell is affected by both solute and pressure potential. The relationship between them is as follows:
Ψw = Ψs + Ψp
11.2.2 Osmosis
The plant cell is surrounded by a cell membrane and a cell wall. The cell
wall is freely permeable to water and substances in solution hence is not
a barrier to movement. In plants the cells usually contain a large central
vacuole, whose contents, the vacuolar sap, contribute to the solute
potential of the cell. In plant cells, the cell membrane and the membrane
of the vacuole, the tonoplast together are important determinants of movement of molecules in or out of the cell. Osmosis is the term used to refer specifically to the diffusion of water across a differentially- or selectively permeable membrane. Osmosis occurs spontaneously in response to a driving force. The net direction and rate of osmosis
depends on both the pressure gradient and concentration gradient. Water
will move from its region of higher chemical potential (or concentration) to its region of lower chemical potential until equilibrium is reached. At equilibrium the two chambers should have nearly the same water potential. If the tuber is placed in water, the water enters the cavity in the
potato tuber containing a concentrated solution of sugar due to osmosis. in which the two chambers, A and B, containing solutions are separated by a semi-permeable membrane.
External pressure can be applied from the upper part of the funnel such that no water diffuses into the funnel through the membrane. 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. Numerically osmotic pressure is equivalent to the osmotic potential, but the sign is opposite.Osmotic pressure is the positive pressure applied, while osmotic potential is negative.
11.2.3 Plasmolysis
The behaviour of the plant cells (or tissues) with regard to water movement depends on the surrounding solution. If the external solution balances the osmotic pressure of the cytoplasm, it is said to be isotonic. If the external solution is more dilute than the cytoplasm, it is hypotonic and if the external solution is more concentrated, it is hypertonic. Cells swell in
hypotonic solutions and shrink in hypertonic ones. Plasmolysis occurs when water moves out of the cell and the cell membrane of a plant cell
shrinks away from its cell wall. This occurs when the cell (or tissue) is placed in a solution that is hypertonic (has more solutes) to the protoplasm. Water moves out; it is first lost from the cytoplasm and then from the vacuole. The water when drawn out of the cell through
diffusion into the extracellular (outside cell) fluid causes the protoplast to
shrink away from the walls. The cell is said to be plasmolysed. The movement of water occurred across the membrane moving from an area of high water potential (i.e., the cell) to an area of lower water potential outside the cell When the cell (or tissue) is placed in an isotonic solution, there is no net flow of water towards the inside or outside. If the external solution balances the osmotic pressure of the cytoplasm it is said to be isotonic. When water flows into the cell and out of the cell and are in equilibrium, the cells are said to be flaccid. The process of plasmolysis is usually reversible. 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. The pressure exerted by the protoplasts due to entry of water against the rigid walls is called pressure potential Ψp.. Because of the rigidity of the cell wall, the cell does not rupture. This turgor pressure is ultimately responsible for enlargement and extension growth of cells. Ψ
11.2.4 Imbibition
Imbibition is a special type of diffusion when water is absorbed by
solids – colloids – causing them to increase in volume. The classical
examples of imbibition are absorption of water by seeds and dry wood.
The pressure that is produced by the swelling of wood had been used by
prehistoric man to split rocks and boulders. If it were not for the pressure
due to imbibition, seedlings would not have been able to emerge out of
the soil into the open; they probably would not have been able to establish!
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. Water potential gradient between the absorbent and the liquid imbibed is essential for imbibition. In addition, for any substance to imbibe any liquid, affinity between the adsorbant and
the liquid is also a pre-requisite.
11.3 LONG DISTANCE TRANSPORT OF WATER
Long distance transport of substances within a plant cannot be by
diffusion alone. Diffusion is a slow process. It can account for only short
distance movement of molecules. For example, the movement of a molecule
across a typical plant cell (about 50 µm) takes approximately 2.5 s.
In large and complex organisms, often substances have to be moved
to long distances. Sometimes the sites of production or absorption and
sites of storage are too far from each other; diffusion or active transport
would not suffice. Special long distance transport systems become
necessary so as to move substances across long distances and at a much
faster rate. Water and minerals, and food are generally moved by a mass
or bulk flow system. Mass flow is the movement of substances in bulk or
en masse from one point to another as a result of pressure differences
between the 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).
The bulk movement of substances through the conducting or vascular
tissues of plants is called translocation. The higher plants have highly specialised vascular tissues – xylem and phloem. Xylem is associated with translocation of mainly water, mineral salts, some organic nitrogen and hormones, from roots to the aerial parts of the plants. The
phloem translocates a variety of organic and inorganic solutes, mainly
from the leaves to other parts of the plants.
11.3.1 How do Plants Absorb Water?
The responsibility of absorption of water and minerals is more specifically
the function of the root hairs that are present in millions at the tips of the
roots. Root hairs are thin-walled slender extensions of root epidermal
cells that greatly increase the surface area for absorption. Water is
absorbed along with mineral solutes, by the root hairs, purely by diffusion.
Once water is absorbed by the root hairs, it can move deeper into root
layers by two distinct pathways:
• apoplast pathway
• symplast pathway
The apoplast is the system of adjacent cell walls that is continuous throughout the plant, except at the casparian strips of the endodermis
in the roots. The apoplastic movement of water occurs exclusively through the intercellular spaces and the walls of the cells. Movement through the apoplast does not involve crossing the cell membrane. This movement is dependent on the gradient. The apoplast does not provide any barrier to water movement and water movement is through mass flow. As water evaporates into the intercellular spaces or the atmosphere, tension develop in the continuous stream of water in the apoplast, hence mass flow of water occurs due to the adhesive and cohesive properties of water. The symplastic system is the system of interconnected protoplasts. Neighbouring cells are connected through cytoplasmic strands that extend through plasmodesmata. During symplastic movement, the water travels through the cells – their cytoplasm; intercellular movement is through the plasmodesmata. Water has to enter the cells through the
cell membrane, hence the movement is relatively slower. Movement is again down a potential gradient. Symplastic movement may be aided by
cytoplasmic streaming. You may have observed cytoplasmic streaming
in cells of the Hydrilla leaf; the movement of chloroplast due to streaming
is easily visible. mMost of the water flow in the roots occurs via the apoplast since the cortical cells are loosely packed, and hence offer no resistance to water movement. However, the inner boundary of the cortex, the endodermis, is impervious to water because of a band of suberised matrix called the 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. Some plants have additional structures associated with them that help in water (and mineral) absorption. A mycorrhiza is a symbiotic association of a fungus with a root system. The fungal filaments form a network around the young root or they penetrate the root cells. The hyphae have a very large surface area that absorb mineral ions and water from the soil from a much larger volume of soil that perhaps a root cannot do. The fungus provides minerals and water to the roots, in turn the roots provide sugars and N-containing compounds to the mycorrhizae. Some plants have an obligate association with the mycorrhizae. For example, Pinus seeds cannot germinate and establish
without the presence of mycorrhizae.
11.3.2.1 Root Pressure
As various ions from the soil are actively transported into the vascular
tissues of the roots, water follows (its potential gradient) and increases
the pressure inside the xylem. This positive pressure is called root
pressure, and can be responsible for pushing up water to small heights
in the stem. Choose a small soft-stemmed plant and on a day, when there is plenty of atmospheric moisture, cut the stem horizontally near the base with a sharp blade, early in the morning.Effects of root pressure is also observable at night and early morning when evaporation is low, and excess water collects in the form of droplets around special openings of veins near the tip of grass blades, and leaves of many herbaceous parts. Such water loss in its liquid phase is known as guttation. Root pressure can, at best, only provide a modest push in the overall process of water transport. They obviously do not play a major role in water movement up tall trees. The greatest contribution of root pressure may be to re-establish the continuous chains of water molecules in the xylem which often break under the enormous tensions created by transpiration. Root pressure does not account for the majority of water transport; most plants meet their need by transpiratory pull.
11.3.2.2 Transpiration pull
Despite the absence of a heart or a circulatory system in plants, the
upward flow of water through the xylem in plants can achieve fairly high
rates, up to 15 metres per hour. Most researchers agree that water is mainly ‘pulled’ through the plant, and that the driving force for this process is transpiration from the leaves. This is referred to as the cohesion-tension-transpiration pull model of water transport. Water is transient in plants. Less than 1 per cent of the water reaching the leaves is used in photosynthesis and plant growth. Most of it is lost through the stomata in the leaves. This water loss is known as transpiration.
11.4 TRANSPIRATION
Transpiration is the evaporative loss of water by plants. It occurs mainly
through stomata (sing. : stoma). Besides the loss of water vapour in
transpiration, exchange of oxygen and carbon dioxide in the leaf also occurs through these stomata. Normally stomata are open in the day time and close during the night. The immediate cause of the opening or closing of stomata is a change in the turgidity of the guard cells. The inner wall of
each guard cell, towards the pore or stomatal aperture, is thick and elastic.
When turgidity increases within the two guard cells flanking each stomatal
aperture or pore, the thin outer walls bulge out and force the inner walls
into a crescent shape. The opening of the stoma is also aided due to the
orientation of the microfibrils in the cell walls of the guard cells. Cellulose
microfibrils are oriented radially rather than longitudinally making it easier for the stoma to open. When the guard cells lose turgor, due to water loss (or water stress) the elastic inner walls regain their original shape, the guard cells become flaccid and the stoma closes. Usually the lower surface of a dorsiventral (often dicotyledonous) leaf has a greater number of stomata while in an isobilateral (often monocotyledonous) leaf they are about equal on both surfaces. Transpiration is affected by several external factors: temperature, light, humidity, wind speed. Plant factors that affect transpiration include number and distribution of stomata, per cent of open
stomata, water status of the plant, canopy structure etc.
The transpiration driven ascent of xylem sap depends mainly on the
following physical properties of water:
• Cohesion – mutual attraction between water molecules.
• Adhesion – attraction of water molecules to polar surfaces (such
as the surface of tracheary elements).
• Surface Tension – water molecules are attracted to each other in
the liquid phase more than to water in the gas phase.
These properties give water high tensile strength, i.e., an ability to
resist a pulling force, and high capillarity, i.e., the ability to rise in thin
tubes. In plants capillarity is aided by the small diameter of the tracheary
elements – the tracheids and vessel elements. The process of photosynthesis requires water. The system of xylem vessels from the root to the leaf vein can supply the needed water. 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. 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’ Measurements reveal that the forces generated by transpiration can create pressures sufficient to lift a xylem sized column of water over 130 metres high.
11.4.1 Transpiration and Photosynthesis – a Compromise
Transpiration has more than one purpose; it
• creates transpiration pull for absorption and transport of plants
• supplies water for photosynthesis
• transports minerals from the soil to all parts of the plant
• cools leaf surfaces, sometimes 10 to 15 degrees, by evaporative
cooling
• maintains the shape and structure of the plants by keeping cells
turgid
An actively photosynthesising plant has an insatiable need for water.
Photosynthesis is limited by available water which can be swiftly depleted
by transpiration. The humidity of rainforests is largely due to this vast
cycling of water from root to leaf to atmosphere and back to the soil.
The evolution of the C4 photosynthetic system is probably one of the
strategies for maximising the availability of CO2 while minimising water
loss. C4 plants are twice as efficient as C3 plants in terms of fixing carbon
dioxide (making sugar). However, a C 4 plant loses only half as much water
as a C3 plant for the same amount of CO2 fixed.
11.5 UPTAKE AND TRANSPORT OF MINERAL NUTRIENTS
Plants obtain their carbon and most of their oxygen from CO2 in the
atmosphere. However, their remaining nutritional requirements are
obtained from water and minerals in the soil.
11.5.1 Uptake of Mineral Ions
Unlike water, all minerals cannot be passively absorbed by the roots.
Two factors account for this: (i) minerals are present in the soil as charged
particles (ions) which cannot move across cell membranes and (ii) the concentration of minerals in the soil is usually lower than the concentration
of minerals in the root. Therefore, most minerals must enter the root by
active absorption into the cytoplasm of epidermal cells. This needs energy
in the form of ATP. The active uptake of ions is partly responsible for the
water potential gradient in roots, and therefore for the uptake of water by
osmosis. Some ions also move into the epidermal cells passively.
Ions are absorbed from the soil by both passive and active transport.
Specific proteins in the membranes of root hair cells actively pump ions
from the soil into the cytoplasms of the epidermal cells. Like all cells, the
endodermal cells have many transport proteins embedded in their plasma
membrane; they let some solutes cross the membrane, but not others.
Transport proteins of endodermal cells are control points, where a plant
adjusts the quantity and types of solutes that reach the xylem. Note
that the root endodermis because of the layer of suberin has the ability to
actively transport ions in one direction only.
11.5.2 Translocation of Mineral Ions
After the ions have reached xylem through active or passive uptake, or a combination of the two, their further transport up the stem to all parts of
the plant is through the transpiration stream. The chief sinks for the mineral elements are the growing regions of the plant, such as the apical and lateral meristems, young leaves, developing flowers, fruits and seeds, and the storage organs. Unloading of mineral ions occurs at the fine vein endings through diffusion and active uptake by these cells.
Mineral ions are frequently remobilised, particularly from older,
senescing parts. Older dying leaves export much of their mineral content
to younger leaves. Similarly, before leaf fall in decidous plants, minerals
are removed to other parts. Elements most readily mobilised are phosphorus, sulphur, nitrogen and potassium. Some elements that are
structural components like calcium are not remobilised.
An analysis of the xylem exudates shows that though some of the
nitrogen travels as inorganic ions, much of it is carried in the organic
form as amino acids and related compounds. Similarly, small amounts
of P and S are carried as organic compounds. In addition, small amount
of exchange of materials does take place between xylem and phloem.
Hence, it is not that we can clearly make a distinction and say categorically
that xylem transports only inorganic nutrients while phloem transports
only organic materials, as was traditionally believed.
11.6 PHLOEM TRANSPORT: FLOW FROM SOURCE TO SINK
Food, primarily sucrose, is transported by the vascular tissue phloem
from a source to a sink. Usually the source is understood to be that
part of the plant which synthesises the food, i.e., the leaf, and sink, the
part that needs or stores the food. But, the source and sink may be
reversed depending on the season, or the plant’s needs. Sugar stored
in roots may be mobilised to become a source of food in the early spring
when the buds of trees, act as sink; they need energy for growth and development of the photosynthetic apparatus. Since the source-sink relationship is variable, the direction of movement in the phloem can
be upwards or downwards, i.e., bi-directional. This contrasts with
that of the xylem where the movement is always unidirectional, i.e.,
upwards. Hence, unlike one-way flow of water in transpiration, food
in phloem sap can be transported in any required direction so long
as there is a source of sugar and a sink able to use, store or remove
the sugar. Phloem sap is mainly water and sucrose, but other sugars, hormones and amino acids are also transported or translocated through phloem.
11.6.1 The Pressure Flow or Mass Flow Hypothesis
The accepted mechanism used for the translocation of sugars from source
to sink is called the pressure flow hypothesis. (see Figure 11.10). As
glucose is prepared at the source (by photosynthesis) it is converted to
sucrose (a dissacharide). The sugar is then moved in the form of sucrose
into the companion cells and then into the living phloem sieve tube cells
by active transport. This process of loading at the source produces a
hypertonic condition in the phloem. Water in the adjacent xylem moves
into the phloem by osmosis. As osmotic pressure builds up the phloem
sap will move to areas of lower pressure. At the sink osmotic pressure
must be reduced. Again active transport is necessary to move the sucrose
out of the phloem sap and into the cells which will use the sugar –
converting it into energy, starch, or cellulose. As sugars are removed, the
osmotic pressure decreases and water moves out of the phloem.
To summarise, the movement of sugars in the phloem begins at the
source, where sugars are loaded (actively transported) into a sieve tube.
Loading of the phloem sets up a water potential gradient that facilitates
the mass movement in the phloem.
Phloem tissue is composed of sieve tube cells, which form long columns
with holes in their end walls called sieve plates. Cytoplasmic strands pass
through the holes in the sieve plates, so forming continuous filaments. As
hydrostatic pressure in the sieve tube of phloem increases, pressure flow
begins, and the sap moves through the phloem. Meanwhile, at the sink,
incoming sugars are actively transported out of the phloem and removed
as complex carbohydrates. The loss of solute produces a high water
potential in the phloem, and water passes out, returning eventually to xylem.
A simple experiment, called girdling, was used to identify the tissues
through which food is transported. On the trunk of a tree a ring of bark
up to a depth of the phloem layer, can be carefully removed. In the absence
of downward movement of food the portion of the bark above the ring on
the stem becomes swollen after a few weeks. This simple experiment
shows that phloem is the tissue responsible for translocation of food; and
that transport takes place in one direction, i.e., towards the roots.
