Transport in Plants

• Explain the need for transport systems in

multicellular plants in terms of size and surface
area:volume ratio;
• Describe, with the aid of diagrams and
photographs, the distribution of xylem and
phloem tissue in roots, stems and leaves of
dicotyledonous plants;
• Describe, with the aid of diagrams and
photographs, the structure and function of xylem
vessels, sieve tube elements and companion
cells;
 Transport in Plants
• Plants need a transport
system so that cells deep
within the plants tissues
can receive the nutrients
they need for cell
processes
• The problem in plants is
that roots can obtain
water, but not sugar, and
leaves can produce
sugar, but can’t get water
from the air
 What substances need to be
moved?
• The transport system
in plants is called
vascular tissue
• Xylem tissue
transports water and
soluble minerals
• Phloem tissue
transports sugars
 The Vascular Tissues
• Xylem and phloem
are found together in
vascular bundles, that
sometimes contain
other tissues that
support and
strengthen them
 Root vs. stem vs. leaf
The vascular bundle
differs depending on if
it is a root or stem
 Root
• The vascular bundle is found
in the centre
• There is a large central core of
xylem- often in an x-shape
• This arrangement provides
strength to withstand the
pulling forces to which roots
are exposed
• Around the vascular bundle
are cells called the endodermis
which help to get water into the
xylem vessels
• Just inside the endodermis is
the periycle which contains
meristem cells that can divide
(for growth)
 Stem
• The vascular bundles are found near the outer edge of the stem
• The xylem is found towards the inside of each vascular bundle, the
phloem is found towards the outside
• In between the xylem and phloem is a layer of cambium
• Cambium is a layer of meristem cells that divide to make new xylem
and phloem
 Leaf
• The vascular bundles
(xylem and phloem) form
the midrib and veins of
the leaf
• A dicotyledon leaf has a
branching network of
veins that get smaller as
they branch away from
the midrib
• Within each vein, the
xylem can be seen on top
of the phloem
A = Xylem
B = Phloem
C/D = Upper/Lower epidermis

Leaf
Xylem vessel wall

Xylem vessel
lumen

Phloem

Endodermis

Starch
grains

Root
Phloem

Xylem

Stem
 Functions of the Xylem
• Provides support for the structure of the
plant
• Transports water and dissolved
substances from root to leaf through
capillary action.
 Structure of the Xylem
• Consists of tubes for water, fibres for
support and living parenchyma cells
• Generally, there are two types of xylem
found in plants; these are:
• (i) Protoxylem
• (ii) Metaxylem.
 Protoxylem
• Protoxylem is the first part of the primary
xylem that matures (Raven et al 1999) and
these cells are usually smaller than the
metaxylem. Metaxylem is the part of the
primary xylem that differentiates after the
protoxylem (Raven et al 1999) and these
cells are usually larger than the
protoxylem.
 Protoxylem
• “Proto” means first. These are
the first vessels formed.
• Protoxylem acts as a
reinforcement for the
metaxylem growth and
development.
• Protoxylem can continue to
elongate as the walls of these
vessels are not completely
lignified.
• The type of lignification found
in Protoxylem is usually
annular or spiral
 Metaxylem
• The part of the primary xylem that
differentiates after the protoxylem and that
is distinguished typically by broader
tracheids and vessels with pitted or
reticulate walls.
 Metaxylem
• These vessels develop after
the protoxylem. Their walls are
entirely lignified after maturity
(more than protoxylem), hence
the cells cannot elongate.
Metaxylem has a reticulate
lignification (spirals are close
together), scalariform and
pitted.
When the Metaxylem cells are
mature, they die and are joined
end to end forming long
narrow tubes which allow the
movement of water.
 Xylem
• NB: Primary xylem develops with the
young plant as it grows (vertically)
• Secondary xylem grows horizontally or
laterally giving a ring-like look to trees.
Trunks of trees become wider due to
the growth of secondary xylem, even
after the plant has completely matured.
 Tissues of the Xylem
• The various tissues
found in the Xylem
are:
• Fibres
• Xylem
Parenchyma
• Tracheids
• Vessels
 Tracheid
• These are single elongated cells with
tapering ends. The tapering end walls
overlap with neighboring tracheid cells.
The tracheids function to give
mechanical support to the plant, they
are also dead and have empty lumens
when they become mature.
Tracheids contain tiny pits in the walls
either in regions that were not lignified
or where the plasmodesmata was
originally. These pits allow for lateral
conduction of water between
tracheids. Water can also move
through the lumen of tracheids that
overlap.
NB: Lignified walls of xylem form
wood.
 Vessels
• Vessels are the primary conducting
unit of the Xylem.
• They are made up of long tubular
structures formed by the fusion of
several cells joined end to end. As
vessels mature, their cell walls
become impregnated with lignin, this
causes them to die (cell contents
break down causing the many cells
that are joined to form a tube.)
Thickening of the cell wall by lignin
occurs in various patterns, some of
which are:
• Ring (Annular)
• Spiral
• Networks
 Xylem Parenchyma
• This is the only living part of the xylem
when it is completely mature.
• It is found in both the secondary and
primary xylem. It allows for the
deposition of tannins (extracted to give
dyes) and crystals.
- It transports food and water along
with carrying out gaseous exchange.
- It maintains a living link through the
pith and the cortex.
• Some of them give rise to xylem
vessels and the phloem tube.
They have meristematic properties;
they divide and differentiate
 Xylem/Sclerenchyma Fibres
• They are similar in
structure to the
tracheid, however
they are shorter,
narrower and thicker
(in terms of wall
lignifications, calls are
dead). They provide
support and they do
not carry water.
Xylem Tissues
 Adaptation of the Xylem for its
Functions
• Cells are long and arranged end to end to form a continuous column.
• Cell contents die when mature, so, there is cytoplasm or nucleus to prevent
water flow.
• Cell walls break down completely; there is no barrier to water flow from
adjacent cells.
• Cell walls are thickened with lignin which makes them more rigid and less
likely to collapse under tension created by the transpiration pull. This also
increases the adhesion of water molecules (water molecules are attracted to
the cell walls because of lignin), enabling them to rise by capillarity.
• Annular matriculate and spiral thickening allow vessels to elongate during
growth and make them more flexible so that the branches can bend in the
wind.
• There are pits throughout the cell to allow lateral movements of water.
• They have a longer lumen of vessels so that a larger volume of water can
be transported.
Movement of Water
and Minerals
through the Plant
 Uptake of Minerals by the Root
Hair Cells
• Roots take up minerals from the soil by ACTIVE TRANSPORT
• (Movement of particles from an area of low concentration to an area of high
concentration)
• Plants need a constant supply of mineral ions to make important molecules such as
chlorophyll and proteins.
• Roots tend have a higher concentration of mineral salts than the soil, therefore when
the mineral ions move from the soil to the root, they move AGAINST a concentration
gradient.
• There is a need of a carrier protein, as the ions cannot cross the membrane freely. This
is due to the fact that the membrane is hydrophobic and the ions are hydrophilic.
Active transport requires ATP, thus respiration of the root hair cells is necessary.
• If respiration is inhibited (e.g. The addition of hydrogen cyanide) the roots will not take
up any minerals because it’s an active process which requires ATP made in the
respiration process.
• Also, some of the minerals are moved across the membrane, via facilitated diffusion,
which is a passive transport method. However the primary mechanism is via active
transport.
 Uptake of Water by Root Hair
Cells
• The root has a low water potential relative
to the soil that surrounds it, because
mineral ions are constantly being taken in.
• Hence, the water will move from the soil
that has a high water potential, into the
root hair cells which have a lower water
potential. This process is known as
osmosis.
 Uptake of Water by Root Hair
Cells
• The root has a low water potential relative
to the soil that surrounds it, because
mineral ions are constantly being taken in.
• Hence, the water will move from the soil
that has a high water potential, into the
root hair cells which have a lower water
potential. This process is known as
osmosis.
Movement of Water Across the
Root
• To move between the tissues in the root
towards the xylem, water takes 3
pathways:
• Apoplast
• Symplast
• Vacuolar
 Water route between cells
• Apoplast: between cell
walls of neighbouring
cells
• Symplast: through
plasma membrane and
plasmodesmata to
cytoplasms from cell to
cell
• Vacuolar: same as
symplast, but also
through vacuoles
 Movement of Water Across the
Root
• Symplast Pathway-water moves through the cytoplasm
via the plasmodesmata. Each cell sets up a concentration gradient
to allow the movement of water particles by osmosis.
 Movement of Water Across the
Root
Vacuolar Pathway-a concentration gradient is set up
and the water molecules move from one cell vacuole to the
next.
 Movement of Water Across the
Root
• Apoplast Pathway-Water moves through the
cell wall. About 50% of the cell wall is empty
space, there the water can move through
unhindered.
 Movement of Water Across the
Root
• Apoplast Pathway
• The cells of the cortex like all plant cells are surrounded by cell walls made of
layers of cellulose fibres crisscrossing one another. Water can soak into these
walls, rather as it would soak into blotting paper and can seep across the root from
cell wall to cell wall without crossing the cytoplasm.
 Movement of Water Across the
Root
• Apoplast Pathway
• Once the water reaches the endodermis,the apoplastic pathway is abruptly
blocked. The cells in the endodermis have a thick, waterproof, waxy band of
suberin in their cell walls. This band, called the Casparian strip, forms an
impenetrable barrier to water in the walls of the endodermis cells. The only
way for water to cross the endodermis is through the cytoplasm of the
endodermal cells.
• As the endodermal cells get older, the suberin deposits become more
extensive, except in certain cells called passage cells, through which water
can continue to pass freely. It is thought that this arrangement gives a plant
control over what mineral ions pass into its xylem vessels, as everything has
to cross cell surface membranes. It may also help with the generation of
root pressure. Once across the endodermis, water continues to move down
the water potential gradient across the pericycle and towards the xylem
vessels.
 Movement of Water Across the
Root
• Apoplast Pathway
• Once the water reaches the endodermis,the apoplastic pathway is abruptly
blocked. The cells in the endodermis have a thick, waterproof, waxy band of
suberin in their cell walls. This band, called the Casparian strip, forms an
impenetrable barrier to water in the walls of the endodermis cells. The only
way for water to cross the endodermis is through the cytoplasm of the
endodermal cells.
• As the endodermal cells get older, the suberin deposits become more
extensive, except in certain cells called passage cells, through which water
can continue to pass freely. It is thought that this arrangement gives a plant
control over what mineral ions pass into its xylem vessels, as everything has
to cross cell surface membranes. It may also help with the generation of
root pressure. Once across the endodermis, water continues to move down
the water potential gradient across the pericycle and towards the xylem
vessels.
 Casparian Strip
• Blocks the apoplast pathway (cell walls)
• Water and dissolved nitrate ions have to pass
into the cell cytoplasm through cell membranes
• There are transporter proteins in the cell
membranes that actively transport nitrate ions
into the xylem lowering the water potential (more
negative)
• Water enters xylem down concentration gradient
and cannot pass back
 Movement across the root
• Active process occurring at the endodermis (layer of cells surrounding the
xylem, some containing waterproof strip called casparian strip)
• Casparian strip blocks the apoplast pathway (between cells) forcing water
into the symplast pathway (through the cytoplasm)
• The endodermis cells move minerals by active transport from the cortex into
the xylem, decreasing the water potential (more negative), thus water moves
from the cortex through the endodermal cells to the xylem by osmosis
• A water potential gradient exists across the whole cortex, so water is moved
along the symplast pathway (through cytoplasm) from the root hair cells
across the cortex and into the xylem
 Water movement up stem
• Root pressure: minerals move into xylem by
active transport, forcing water into xylem and
pushes it up the stem
• Transpiration Pull: loss of water at leaves
replaced by water moving up xylem. Cohesion-
tension theory- cohesion between water
molecules and tension in the column of water
(which is why xylem is strengthened with lignin)
means the whole column of water is pulled up in
one chain
• Capillary action: adhesion of water to xylem
vessels as they are narrow
 How does water move up the
xylem?
• Water can move through the xylem parenchyma via
osmosis, but they cannot move through the xylem
vessels and the tracheids as they are dead so they have
no cytoplasm etc.
• There are three methods by which water can move up
the xylem; all three of these methods work together.
• Capillarity (cohesion-tension)
• Root pressure
• Transpiration pull
 Capillarity
• Capillarity – the movement of fluid up a narrow tube.
•
Cohesive forces between like molecules and adhesive forces between unlike
molecules facilitate capillarity. The water molecules in the xylem are said to be under
tension as one water molecules pull others along with it up the xylem.
• Capillarity helps but it doesn’t account for the movement of water up the tube alone.
There are other things at play.
 Root Pressure
• Water is constantly moving into the plant
roots via osmosis. This creates root
pressure that drives the water up the
xylem.
 Root Pressure
• Root pressure sometimes results in
guttation, the exudation of water droplets
on tips or edges of leaves … usually in
small plants
 Transpiration Pull
• Transpiration pull or the suction force is the force which aids in drawing the
water upward from roots to leaves. In leaves, some amount of water is used for
photosynthesis and excess water is released into atmosphere through openings
called as stomata
• Transpiration is the evaporation of water from plant surfaces. When the
water gets to the leaf, it eventually leaves the leaves via diffusion
through the stomata.
As a result low pressure is created at the top of the xylem relative to the
root xylem where water constantly rushes in resulting in a high pressure.
A pressure gradient is set up. This creates the transpiration pull.

Water evaporates from the leaves, through the stomata. As this

happens, water is pulled up from the xylem, into the leaf to replace that,
which has been lost, this is known as the transpiration pull. The
transpiration pulls puts the xylem under tension.
 Cohesion-Tension Theory
• The 'Cohesion-Adhesion Tension' theory explains the translocation
of water from the root system to the shoot system has been
proposed by Dixon and Joly (1894). According to them following
factors help the ascent of sap (water and mineral ions).
1. Strong cohesive-adhesive forces of water among the own
molecules and the wall of xylem help to maintain continuity of water
in the vascular bundles.
2. Presence of continuous water column from the root to the shoot
of the plants.
3. The transpiration generates a transpiration pull from the leaves of
the plants to roots through the vascular bundles.
Cohesion-Tension Theory
Cohesion-Tension Theory
Cohesion-Tension Theory
Cohesion-Tension Theory
 Evidence to prove Cohesion-
tension
• The change, which occurs in the diameter of the leaves
according to the rate of transpiration. The diameter of the tree
trunk or stem is smaller in the day than in the night.
- The walls of the Xylem are being pulled inwards as they are under
more tension. At night when transpiration is at its lowest there is
less tension in the Xylem and so the diameter increases.

• When a Xylem vessel is broken, water does not leak out which
would be the case if it were under pressure, but rather air is
pulled in which is consistent with it being under tension
(Suction Pressure).
Physical Challenges of
Cohesion-Tension
Minimising the Effect of
Cavitation
Minimising the Effect of
Cavitation
 Transpiration: three processes
• Osmosis from xylem to mesophyll cells
• Evaporation from surface of mesophyll
cells into intercellular spaces
• Diffusion of water vapour from intercellular
spaces out through stomata
 Transpiration
• Loss of water vapour from upper parts of the
plant
• Water enters leaf from xylem and passes to
mesophyll cells by osmosis
• Water evaporates from surface of mesophyll
cells to form water vapour (air spaces allow
water vapour to diffuse through leaf tissue)
• Water vapour potential rises in air spaces, so
water molecules diffuse out of the leaf through
open stomata
 How does water move through
the leaves
• Water is constantly being evaporated from the cells via the stoma, there is always a
concentration gradient is that set up with the cells in relation to the xylem. The cells of the leaf
have a low water potential while the xylem has a high water potential. The water will diffuse from
the xylem into the spongy mesophyll or palisade mesophyll cell.
The water moves to surrounding cells via three pathways:
1. Apoplast
• 2. Symplast
3.Vacuolar

As the water moves from the palisade cell, it enters the cells of the upper epidermis and some is
lost through the cuticle.
The water that moves into the spongy mesophyll cell, some of it is diffuses into the sub-stomatal
air space. When there is an accumulation of water vapour in the sub-stomatal air space, it then
moves through the stomata into the atmosphere.
• NB: The leaf loses water in two ways, wither through the epidermis or the stomata, however, due
to the presence of a waxy cuticle above the upper epidermis only small amounts of water is lost
through it. As a result 90% of water is lost through the stomata.
 How water leaves the leaf
• Through stomata
• Tiny amount through the waxy
cuticle
• Water evaporates from the
cells lining the cavity between
the guard cells, lowering water
potential and meaning that
water enters them by osmosis
from neighbouring cells, which
is replaced by further
neighbouring cells and so on
 Water use in plant
• Photosynthesis
• Cell growth and elongation
• Turgidity
• Carriage of minerals
• Cools the plant
 Transpiration
• 97-99.5% of the water drawn up by plants
is lost due to transpiration.
• Why is transpiration important?
Importance of Transpiration
 Measuring transpiration
• Potometer is used to
estimate water loss
 Factors affecting transpiration
• Leaf number: more leaves, more transpiration
• Number, size, position of stomata: more and large, more
transpiration, under leaf, less transpiration
• Cuticle: waxy cuticle, less evaporation from leaf surface
• Light: more gas exchange as stomata are open
• Temperature: high temperature, more evaporation, more
diffusion as more kinetic energy, decrease humidity so
more diffusion out of leaf
• Humidity: high humidity, less transpiration
• Wind: more wind, more transpiration
• Water availability: less water in soil, less transpiration
(e.g. in winter, plants lose leaves)
 External Factors
• External Factors That Affect the Rate of Transpiration

1. TEMPERATURE

The higher the temperature, the kinetic energy the water molecules have, and so they are able to
break the attractive forces between them which will result in evaporation happening more quickly.
As they evaporate, they move out of the plant.
•
- High temperature speeds of transpiration by providing the latent heat of vapourization, at the
same time, high temperature also lowers the relative humidity of the air; both of these increase
the gradient in the water vapour concentration between the sub-stomatal cavity and the external
atmosphere.
•
• 2. LIGHT
• The stomata tends to be open and as water vapour forms it tends to diffuse through the stomata.
This is due to the fact that it facilitates the light-independent stage of photosynthesis.

- High light intensities stimulate the opening of the stomata which results in the transpiration rate
increasing. This is closely linked to high temperatures. If the rate of transpiration is too high, the
stomata will close.
 External Factors
• External Factors That Affect the Rate of Transpiration

3. AIR MOVEMENT
• Moving air sweeps away the water vapour in the air outside the stomata speeding up diffusion of water vapour by making a steeper gradient between the sub-stomatal
cavity and the air. Consequently, windy conditions lead to an increased rate of transpiration.
•
• 4. HUMIDTY
• High humidity decreases the rate of transpiration, by making the concentration gradient less steep.
•
• 5. ATMOSPHERIC PRESSURE
Water vapour pressure decreases as the atmospheric pressure decreases. The lower the atmospheric pressure, the greater the rate of evaporation form the leaves. The
pressure of the atmosphere ultimately keeps the water vapour from evaporating, therefore if it is low, then it allows for more water to evaporation.

•
• 6. WATER SUPPLY
• A shortage in water supply leads to wilting, and when the plant begins to wilt, the stomata will close. Therefore the lower the water supply, the slower the rate of
transpiration.
 Internal Factors
• Internal Factors That Affect the Rate of Transpiration
•
• 1. THE LEAF SURFACE AREA
• The greater the surface area the greater the rate of transpiration; more of the leaf is exposed
hence the more water will be lost.
- For plants in the dessert they have thick leaves or cells
•
• 2. THICKNESS OF CUTICLE
The thinner the cuticle, the higher the rate of transpiration. The cuticle designed to is to prevent
water loss.

3. NUMBER OF STOMATA
The more stomata the leaves have, the greater the rate of transpiration.
•
• 4. DISTRIBUTION OF THE STOMATA
If more stomata are in the upper epidermis, the greater the rate of transpiration; this is due to
direct exposure to the sun.
Opening and Closing of the
Stomata
 Opening and Closing the
Stomata
• i. Stomata open when H+ ions are transported out of the guard cell using ATP
ii. This leads to the inside of the guard cell becoming more negative
iii. As a result K+ ions diffuse into the guard cell, which reduces the water potential of
the guard cells, and water moves in by osmosis.
iv. The volume of the guard cells therefore increases causing them to curve apart due
to then having inner cell walls that are thicker than the outer wall.
 Too much water loss
• Less turgidity
• Non-woody plants wilt and die
• Leaves of woody plants die first then it will
die if water loss continues
 Xerophytes
• Smaller leaves reducing surface area e.g. pine tree
• Densely packed spongy mesophyll to reduce surface area, so less
water evaporating into air spaces
• Thick waxy cuticle e.g. holly leaves to reduce evaporation
• Closing stomata when water availability is low
• Hairs on surface of leaf to trap layer of air close to surface which
can become saturated with water, reducing diffusion
• Pits containing stomata become saturated with water vapour
reducing diffusion
• Rolling the leaves so lower epidermis not exposed to atmosphere
also traps air which becomes saturated
• Maintain high salt concentration to keep water potential low and
prevent water leaving
 Marram Grass
Leaf rolled up to trap
air inside

Thick waxy cuticle to

reduce water
evaporation from the
surface

Trapped air in the

centre with a high
water potential (less
negative)

Hairs on lower
Stomata in pits to surface reduce
trap air with moisture movement of air
close to the stomata
 Structure of Phloem
• Function to transport
products of
photosynthesis from
one part to another
• The Phloem is made up of 5
types of tissues, they are as
follows:
• Sieve Tube Element
• Companion Cells
• Parenchyma
• Fibres
• Sclereids
 Sieve Tubes
• Sieve tube, in flowering plants, elongated
living cells (sieve-tube elements) of the
phloem, the nuclei of which have
fragmented and disappeared and the
transverse end walls of which are pierced
by sieve like groups of pores (sieve
plates). They are the conduits of food
(mostly sugar) transport.
 Sieve Tube Element
• Sieve Tube Element
These are elongated structures that are
joined end to end to form long tubes.
• The cells are living and retain a thin layer of
cytoplasm within their cell surface
membrane, which lies against the cellulose
cell wall.
• Within the cytoplasm are mitochondria and
a modified form of endoplasmic reticulum.
However unlike most cells, there is no
nucleus, Golgi apparatus, or ribosomes.
• These structures are broken down in order
to make the sieve tubes more hollow and so
reduce the resistance to flow of liquid within
them
 Sieve Tube Element
• Sieve Tube Element
The end walls of the sieve tubes are
perforated by large pores; these
perforated end walls are called sieve
plates.
• It is through the pores created by the
sieve plates that fluid; containing the
products of photosynthesis as well as
mineral ions, pass from one sieve tube
element to another.

• The space inside the sieve tube

element is known as a lumen.
• NB: The sieve tube element is a
living structure unlike the vessel
element in the xylem.
 Companion cells
• In between sieve tubes
• Large nucleus, dense
cytoplasm
• Many mitochondria to
load sucrose into sieve
tubes
• Many plasmodesmata
(gaps in cell walls
between companion cells
and sieve tubes) for flow
of minerals
 Companion Cell
• Companion Cells
These are closely associated with
the sieve tube elements, both
come from the same cell division.
Because the sieve tube element
lacks organelles such as nucleus,
ribosomes and Golgi apparatus, it
is unable to perform all the
metabolic process that are
essential for its survival. The
companion cell is the sight of all
these processes; in other words,
the companion cell carries out the
metabolic processes for the sieve
tub element as well itself.
 Companion Cell
• Companion Cells
They are both closely linked by
plasmodesmata; so close is the relationship
that if one cell dies so does the other. Some
companion cells have very folded cell walls
and cell surface membranes, which create a
large surface area, to increase the rate of
transfer of photosynthetic products into the
sieve tube elements.
• NB: A function of the companion cell is
to transfer the products of
photosynthesis into the sieve tube
element. Another is to carry out the
metabolic process for the sieve tube
element.
A transfer cell is a specialized companion
cell, however the difference is that they
have in-foldings which increase the surface
area of absorption of sucrose. The function
of the transfer cells is to pump products into
the sieve tubes.
 Phloem Parenchyma
• Phloem
Parenchyma
Similar to those in
the xylem and
carry out the same
function; which is
primarily for food
storage
 Fibres
• Fibres
Same type of fibres
found in xylem,
elongated cells with
tapered ends, they
are similar to the
tracheid but more
narrow and shorter.
Their function is for
support.
 Phloem Sclereids
• Sclereids
They are spherical
and have lignified
walls. Their main
function is to confer
rigidity and firmness;
the Sclereids have a
structural support
role.
 Conducting Elements of the
Phloem
• The phloem has 2 conducting elements
(actively involved in translocation). These
are:
• 1. The Sieve Tube Element
2. Phloem Parenchyma
How is the Phloem Adapeted for
its Function
• Sieve tube elements are elongated and arranged end to end to form a continuous
column.
• The nucleus and many of the organelles are located in the companion cells leaving
the lumen of the sieve tube elements more open so reducing resistance to the flow of
liquid
• Sieve plates are perforated with sieve pores, reducing resistance to liquid flow.
• Sieve plates hold the walls of sieve tube elements together and prevent them from
bursting.
• The walls contain cellulose microfibrils that run around the cell giving strength and
preventing the tube from bursting under pressure.
• The walls are thin to allow easy entry of water at the source, which helps to build up
pressure.
• Companion cells have many mitochondria to release the ATP needed for
translocation of organic materials.
• Plasmodesmata allow easy movement of substances to and from companion cells.
Substances Transported in
phloem sap
Transport in the Phloem-Ernst
Munch Mass Flow Experiment
Mass/Pressure Flow Hypothesis
• Translocation is movement of organic solutes from a source ( area of
manufacture or release of organic solutes-leaves, roots) to a sink. A sink;
can be any part of the plant that receives solutes from the leaves and use
these solutes for metabolic process for example roots, fruits or seeds.
• The products of photosynthesis (coming from the leaf) are constantly being
pumped into the phloem and so it has a low water potential. This results in
water entering the phloem form the xylem, causing a build up of hydrostatic
pressure in the phloem, since water is being constantly being pulled in. This
forces all the solutes in the phloem down to the roots. Once in the roots,
they are used up, converted to starch or compounds that do not affect the
concentration of water in the roots.
The concentration never goes to equilibrium because as the solutes are
being transported to the roots they are used up or converted to insoluble
compounds, so it does not affect the osmotic balance and the roots remains
dilute in comparison to the phloem.
Mass/Pressure Flow Hypothesis
Surprisingly, the exact mechanism of sugar transport in the phloem is not
known, but it is certainly far too fast to be simple diffusion. The main
mechanism is thought to be the mass flow of fluid up the xylem and down the
phloem, carrying dissolved solutes with it.
Plants do not have hearts, so the mass flow is driven by a combination of active
transport and evaporation. This is called Mass Flow Theory and it works like
this:
Mass/Pressure Flow Hypothesis
 Mass/ Pressure Flow
Hypothesis
• Sucrose produced by photosynthesis is actively pumped into the phloem vessels by
the companion cell.
• This decreases water potential in the leaf phloem, so water diffuses from xylem to
xylem by osmosis.
• This increases the hydrostatic pressure in the phloem, so water and dissolved solutes
are forced downwards to relieve the pressure. This is mass flow: the flow of water
together with dissolved solutes due to a force.
• In the roots, the solutes are removed from the phloem by active transport in the cells
of the root.
• At the same time, ions are being pumped into the xylem from soil by active transport
reducing the water potential in the xylem.
• The xylem now has lower water potential than the phloem so water diffuses by
osmosis form the phloem to the xylem.
• Water and its dissolved ions are pulled up the xylem by tension from the leaves. This
is also mass flow.
 Movement of Sugars
• Translocation: movement of assimilates (sugars
and other chemicals) through the plant
• Source: a part of the plant that releases sucrose
to the phloem e.g. leaf
• Sink: a part of the plant
that removes sucrose from
the phloem e.g. root
 Sucrose Entering the Phloem
(Phloem loading)
• Active process (requires energy)
• Companion cells use ATP to transport hydrogen
ions out of their cytoplasm
• As hydrogen ions are now at a high
concentration outside the companion cells, they
are brought back in by diffusion through special
co-transporter proteins, which also bring the
sucrose in at the same time
• As the concentration of sucrose builds up inside
the companion cells, they diffuse into the sieve
tubes through the plasmodesmata (gaps
between sieve tubes and companion cell walls)
Phloem Loading
Sucrose movement through phloem
• Sucrose entering sieve tube lowers the water
potential (more negative) so water moves in by
osmosis, increasing the hydrostatic pressure
(fluid pushing against the walls) at the source
• Sucrose used by cells surrounding phloem and
are moved by active transport or diffusion from
the sieve tube to the cells. This increases water
potential in the sieve tube (makes it less
negative) so water moves out by osmosis which
lowers the hydrostatic pressure at the sink
 Movement along the phloem
• Water entering the phloem at the source,
moving down the hydrostatic pressure
gradient and leaving at the sink produces
a flow of water along the phloem that
carries sucrose and other assimilates. This
is called mass flow. It can occur either up
or down the plant at the same time in
different phloem tubes
 Phloem Unloading
• Phloem Unloading:
• It occurs in the consumption end or sinks organs
(such as developing roots, tubers, reproductive
structures etc.)
• Sugars move from sieve tubes to receiver
cells in the sink in­volving following steps:
• (i) Sieve element unloading:
• In this process, sugars (imported from the
source) leave sieve elements of sink tissues.
 Phloem Unloading
• (ii) Short distance transport:
• The sugars are now transported to cells in sink by a short distance pathway
which has also been called as post-sieve element transport.
• (iii) Storage and metabolism:
• Finally, sugars are stored or metabolized in the cells of the sink.
• As with the phloem loading process, sucrose unloading also occurs through
symplast via plasmodesmata or through apoplast at some point en route to
sink cells.
• Phloem unloading is typically symplastic in growing and respiring sinks such
as meristems roots, and young leaves etc. in which sucrose can be rapidly
metabolized. (Young leaves act as sink until their photosynthetic machinery
is fully developed, at which point they become sources).
 Evidence for translocation
• Radioactively labelled carbon from carbon dioxide can appear in the
phloem
• Ringing a tree (removing a ring of bark) results in sugars collecting
above the ring
• An aphid feeding on the plant stem contains many sugars when
dissected
• Companion cells have many mitochondria
• Translocation is stopped when a metabolic poison is added that
inhibits ATP
• pH of companion cells is higher than
that of surrounding cells
• Concentration of sucrose is higher at
the source than the sink
 Evidence to Prove
Translocation
• When the phloem is cut, sap is seen. When tested organic
molecules will be found.
• When the plant is exposed to radioactive to carbon dioxide and then
it is allowed to photosynthesize. When testing for radioactivity, it is
detected specifically in the phloem.
• The aphids feed on sugars coming from the plant; the sap that
comes from the stylets is tested and organic molecules are found to
be in it.
• If a ring of bark is removed from a tree, and the tree left for a week.
You will notice that the top of the trunk swells up as the solutes
cannot be transferred to the roots (ringing experiment).
 Evidence to Prove the Mass
Flow Theory
• High pressure in the sieve tubes as shown by sap being released when they are cut.
This is in contrast to the xylem as it sucks in air (due to suction pressure)
• The concentration of sucrose in the phloem is about 10%-30%, while the
concentration of sucrose in the leaves is 0.5%.
• The pH of the sieve tubes is about 7.5-8.0 (slightly alkaline); to support the fact that
hydrogen ions are being constantly pumped out of the phloem.
• The concentration of sucrose is higher in the leaf phloem (source) than in the root
phloem (sink).
• Metabolic poisons or a lack of oxygen inhibits translocation of sucrose in the phloem;
this is because the companion cells but respire to produce ATP to pump the
hydrogen ions into the mesophyll cell.
• Companion cells possess many mitochondria and readily produce ATP; it has more
mitochondria than most cells in the plants.
• Downward flow in the phloem occurs in daylight but ceases when leaves are shaded
at night. This is because the plant photosynthesizes in the day.
• Increases in sucrose levels in the leaves are followed by increases in the sucrose
level in the phloem a little later.
Evidence against the Mass Flow
Hypothesis
• The sieve plates serve as an impediment of sap that is flowing
through the phloem. But it is suggesting that it has a structural
function, which helps to prevent the tubes from bursting under
pressure by restricting how much of the sap is travelling through the
phloem at a time.
NB: The sieve plate produce phloem proteins which act a
“band aid” blocking the sieve pores when the phloem is
broken; refer to the ringing experiment.

• Not all the solutes move at the same speed; sucrose might move
faster than the amino acid.
• Does not account for the movement of solute in both directions.
eg. When its spring, leaves grow back. For the leaf to grow back
they need food which comes from the roots.
 Useful Revision Sites
• http://scienceaid.co.uk/
• http://www.s-cool.co.uk/
• http://www.sparknotes.com/biology/