 The transport of organic solutes from

one place to another in higher plants

is referred to as translocation of
organic solutes.

 Translocation of organic solutes

always takes place from the region of
higher concentration of soluble form
to the region of lower concentration of
its soluble form i.e. Supply end
(source) Consumption end
(sink)
 The two long distance transport
pathways, the phloem and the
xylem, extend throughout the plant
body.

 Water is pulled from the roots to the

leaves through a process called Transpiration
transpiration.

 In addition, water potential drives

the movement of water from one
area of the plant to another using
osmosis, gravity, and the surface
tension of the water.
 The xylem is the tissue that transports water and minerals from
the root system to the aerial portions of the plant.

 The phloem is the tissue that translocates the products of

photosynthesis from mature leaves to areas of growth and
storage, including the roots.

 As we will see, phloem also redistributes water and various

compounds throughout the plant body.

Sucrose

Water
 In leafs the spongy mesophyll layer contains
arrangements of vascular tissue, consisting of xylem
and phloem that are specialized for the transport of
water and nutrients throughout the plant.

Mesophyll cells

Phloem

Xylem
 The vascular tissue extends from
leaves(where the water potential is
lower), through the stem to the roots
( where the water potential is
higher).

 The arrangement of the tissue, the

functions of the cells, and water
potential determine the direction in
which water move through a plant.
Water posses out of the leaf as water
vapour through the stomata.

 The water vapour lost from the leaves

is replaced with water that enters
through the roots and is brought up
through the stem in xylem.
 Xylem is composed of vessels,
which are continuous tubes formed
from dead, hollow, cylindrical cells
arranged end to end, and tracheids,
which are dead cells that taper as the
ends of overlap.

 This arrangement and the pore

nature of water molecules allow
water pass in an unbroken stream
through the xylem from the roots up
through the shoots into the leaves.
 Cohesion
 Adhesion is the attraction of
water molecules to a surface, such
as the wall of the xylem.

 Cohesion is hydrogen bonding

between water molecules.

 Together, adhesion and cohesion

allow water to move through the Adhesion
xylem in a continuous stream
from the roots up through the
stem to replace water lost from the
leaves through the stomata.

 Water enters the plant through the

epidermal cells of the roots and
travels into the xylem.
 Symporter pump

 Water potential in the cells of the

roots increases when symporter
pumps in the plasma membrane
allow proton to pass into the cells,
travelling down their concentration
gradient.

 These pumps couple the transport of

protons with the transport of
minerals and other solutes into the
cell, water follows into the cell driven
by osmosis.

 The presence of aquaporin

channels in the membrane enhances
osmosis allowing bulk flow of water
from the soil into the roots.

Aquaporin channel
 In vascular plants, phloem is a complex living tissue
that carries organic nutrients, particularly sucrose,
amino acids, fatty acids, certain signaling factors
to all parts of the plant where needed.

 Phloem tissue consists of less specialized and nucleate

parenchyma cells( in addition to fibres and sclereids).
 Conduction in phloem is carried
out through two kinds of elongated
cells, sieve cells and sieve-tube
members, both types of cells have
clusters of pores known as sieve
areas.

 These structures aid in the

movement of carbohydrates, like
sucrose, that are manufactured in
the leaves and carried in the
phloem throughout the plant, a
process called translocation.
 Turgor pressure increases in the sieve tube members as
sucrose from surrounding cells is brought into
phloem through active transport. Water then enters
phloem from xylem by osmosis which drives the
transport of carbohydrates in the phloem.
 Water movement in vessels is one
way, while transport in sieve tube
members can go in both directions.

 Water potential is important driver in

both xylem and phloem transport, but
only phloem transport utilizes both the
active and passive transport.

 Our heart pumps blood throughout

our bodies to provide nutrients and
water to our cells. Vascular plants can
accomplish this same feature without a
heart, using transpiration, water
potential, and translocation to move
water nutrients, and mineral to all cells
of the plant.
 The best supported theory to explain the movement of food
through the phloem is called the Pressure-Flow Hypothesis
(PFH), also known as Mass Flow Hypothesis.

 This hypothesis was proposed by Ernst Munch, a german

plant physiologist in 1930.

 The pressure flow mechanism depends upon;

 Turgor pressure
 Difference of osmotic pressure gradient along the direction of
flow between the source and sink.
 The Munch hypothesis
postulated the movement of
protoplasm en mass along a
turgor pressure gradient,
induced by maintained
gradient of water potential.

 The mass flow of organic

solutes takes place from the
site of higher
concentration to the site of
lower concentration
The principles of mass flow hypothesis is illustrated in the model shown in
figure
 The two chambers A & B with semi permeable membrane are connected
by a tube T, and reservoirs are filled with water.
 The chamber A contains concentrated sugar solution and chamber B
contains dilute sugar solution.
 The OP of chamber A is high as compared to B. The water enters into
chamber A and its turgor pressure is increased.
 This increase in turgor pressure causes mass flow of sugar solution to
chamber B under the influence of turgor pressure gradient.
 The movement of solute will continue till the solution in both the
chambers attain the same concentration.
 Thus, it is the pressure gradient between source
(leaves) and sink (shoot and roots) that drives the
contents of the phloem up and down through the sieve
tubes.
There are different pieces of evidences
that support the Munch
hypothesis;
 When the woody or herbaceous
plant is girdled, the sap contains
high sugar content exudates from the
cut end.

 Positive concentration gradient

disappears when the plants are
defoliated.

 Movement of viruses and growth

hormones is fast in illuminated
leaves as compared to shaded leaves.
Opposition or criticisms against the
Munch hypothesis are often voiced.
 The hypothesis fails to explain
bidirectional movement of
metabolites Which is common in
plants, i.e. movement of different
substances in opponent directions at
the same time.

 Munch hypothesis gives a passive role

to the sieve tube and protoplasm,
while some workers demonstrated
involvement of ATP.

 Turgor pressure may not always be

higher at the supply end.
 Phloem is transported and loaded
into sieve tube members.

 Phloem loading is the movement

of photosynthetic products from
the mesophyll chloroplasts to the
sieve elements of mature leaves.

 Phloem loading is specific and

selective.
 The movement of sugars from
mesophyll cells to sieve tubes of
phloem may occur either
through symplast or sugars may
enters through apoplast at
same point to phloem sieve
tubes.

1. Apoplastic loading
2. Symplastic loading
 Sugar moves through plasmodesmata
from mesophyll cells up to companion
cells where it moves into the apoplast
and is actively loaded into ordinary
companion cells.

 Ordinary companion cells with cell

wall in growths have very few
plasmodesmata and load from the
apoplast.

 In the apoplastic pathway, sucrose

uptake requires metabolic energy.
 Symplastic phloem loading
dependent on plant species with
intermediary companion cells.

 These requires the presence of

open plasmodesmata between
different cells in the pathway.

 In symplastic phloem loading

type of sugar transported are
sucrose and other
oligosaccherides.
 FROM CHLOROPLASTS TO
SIEVE ELEMENTS –

I. During the day - Triose

phosphate formed by
photosynthesis during the
day is transported from the
chloroplast to the cytosol,
where it is converted to
sucrose.
During the night - carbon
from stored starch exits the
chloroplast probably in the
form of glucose and is
converted to sucrose.
.
2. Sucrose moves from the mesophyll cell to the
vicinity of the sieve elements in the smallest veins of
the leaf.

3. Sieve element loading-

 Sugars are transported into the sieve elements and
companion cells, where they become more
concentrated than in the mesophyll.
 Translocation through the vascular system to the sink
is referred to as long – distance transport.
.
 Sieve Element Loading Involves a
Sucrose –
H+ Symporter---
 A sucrose–H+ symporter is
throught to mediate the transport
of sucrose from the apoplast into
the sieve element–companion cell
complex.
 The sugars are actively loaded from
the apoplast into the sieve
elements and companion cells by
an energy-driven.
 selective transporter located in the
plasma membranes of these cells .
Apoplastic phloem loading leads to
three basic predictions ;
1. Transported sugars should be found in the apoplast .
2. sugars are supplied to the apoplast, the exogenously
supplied sugars should accumulate in sieve elements
and companion cells .
3. Inhibition of sugar uptake from the apoplast should
result in inhibition of export from the leaf.
During phloem unloading following steps occur;
1. Sieve- element unloading – This is the process by
which imported sugars leave the sieve elements of sink
tissues.
2. Short– distance transport – After sieve element
unloading, the sugars are transported to cells in the sink
by means of short distance transport pathway.
 This pathway has been also called post – sieve element
transport.
3. Storage and metabolism – In the final step, sugars are
stored or metabolized in the sink
 In some young dicot leaves, such as sugar beet and tobacco .

 Symplastic pathway of unloading includes insensitivity to PCMBS ( p-

chloromercuribenzenesulfonic acid ), a reagent that inhibits the
transport of sucrose across plasma membranes but does not permeate
the symplastic pathway.

 Meristematic and elongating regions of primary root tips also appear

to unload symplastically. Sufficient plasmodesmata exist in these
pathways to support symplastic unloading.
 Type 1: This phloem unloading pathway is designated
 apoplastic because one step, transport from the sieve
element–companion cell complex to the successive sink
cells, occurs in the apoplast.

 The sugars are taken into the symplast of adjoining cells,

transport is symplastic. This route has not yet been
demonstrated in any sink type.

 Type 2: This pathway also has an apoplastic step.

 However, the exit from the sieve element companion cell
complex i ,e, sieve element unloading is symplastic.

 The apoplastic step occurs - Figure (2A) shows an

apoplastic step close to the sieve element–companion cell
complex;
 Figure (2B), an apoplastic step that is further removed .
 Phloem translocation – Wikipedia
the free encyclopedia

 Phloem loading and unloading –

Wikipedia the free encyclopedia

 Plant physiology – LincolnTaiz

and Eduardo Zieger

 Plant physiology – Frank B.

Salisbury and Cleon W. Ross