Translocation in

the Phloem

Fig. 10.1

Phloem

xylem
The phloem contains a very concentrated
solution of dissolved solutes, mainly sucrose,
but also other sugars, amino acids, and other
metabolites. This solution is called the sap, and
the transport of solutes in the phloem is called
translocation.
 Phloem Structure
The main components of phloem are
sieve elements
companion cells.
Sieve elements have no nucleus and only
a sparse collection of other organelles .
Companion cell provides energy
so-named because end walls are
perforated - allows cytoplasmic
connections between vertically-stacked
cells .
conducts sugars and amino acids - from
the leaves, to the rest of the plant
 Phloem transport requires
specialized, living cells
Cells called sieve tube
elements join to form a
continuous tube

Sieve tube elements lack some

structures and organelles - no
nuclei, vacuole, Golgi,
ribosomes, or microtubules

Pores in sieve plate between

sieve tube elements are open
channels for transport
 Phloem transport requires
specialized, living cells
Each sieve tube element is
associated with one or
more companion cells.
 Phloem transport requires
specialized, living cells
Companion cells:
Transport products of
photosynthesis from cells in
leaves to sieve tube elements
through plasmodesmata

Synthesize the various proteins

used in the phloem

Contain many, many

mitochondria for cellular
respiration to provide the
cellular energy required for
active transport

Cell wall between
sieve elements

Sieve plate pore

Companion cell
 Exactly what
is transported
in phloem?
The phloem is the vascular system for moving (translocating)
sugars produced in photosynthesis and other
substances throughout the plant.

 Sugars in the phloem
Carbohydrates transported in
phloem are all nonreducing
sugars.

Reducing sugars, such as

glucose are too chemically
reactive to be transported in the
phloem

The most common transported

sugar is sucrose.
A disaccharide made up from
glucose & fructose
The mechanism of
phloem transport
The Pressure-Flow Model
 Phloem transports sugars
from a source to a
sink
Source:
Any exporting region that produces sugars above
and beyond that of its own needs
Sink:
Any area that does not produce enough sugar to
meets its own needs
 During the summer sugar is mostly transported from the leaves,
where it is made by photosynthesis (the source) to the roots,
where it is stored (the sink).
During the spring, sugar is often transported from the
underground root store (the source) to the growing leaf buds
(the sink).
Flowers and young buds are not photosynthetic, so sugars can
also be transported from leaves or roots (the source) to flowers
or buds (sinks).
In source tissue
sugars are moved from photosynthetic cells and actively

loaded (uses ATP energy) into companion cells and sieve
tube elements.

Phloem loading uses a
proton/sucrose co-transport prote
in.
 The Pressure -Flow Model
Phloemloadingleadstoa
buildupofsugars(the
phloemcellsbecome
hypertonic)

Inresponse,waterenters
sieveelementsfromxylem
viaosmosis

Thusphloemturgor
pressureincreases
 The Pressure -Flow Model
In sink tissue
Phloemunloadingleadsto
lowersugarconcentration
(thephloemcellsbecome
hypotonic)

Waterleavesthephloem
andenterssinksieve
elementsandxylem(via
osmosis)

Thusphloemturgor
pressuredecreases
Phloem solution moves along a gradient of pressure
generated by a solute concentration difference
between source and sink ends of the pathway

Fig. 10.10
 Massflowhypothesis
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 dont have hearts, so
the mass flow is driven by a combination of active
transport (energy from ATP) and evaporation (energy
from the sun). This is called the mass flow theory,
and it works like this:
1. Sucrose produced by photosynthesis is actively
pumped into the phloem vessels by the companion
cells.

this:

2. This decreases the water potential in the leaf phloem,

so water diffuses from the neighbouring xylem
vessels by osmosis.
3. This is 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
its dissolved solutes due to a force.
4. In the roots the solutes are removed from
the phloem by active transport into the cells of
the root.

5. At the same time, ions are being pumped

into the xylem from the soil by active
transport, reducing the water potential in the
6. The xylem now has a lower water potential
than the phloem, so water diffuses by osmosis
from the phloem to the xylem.
7. Water and its dissolved ions are pulled up
the xylem by tension from the leaves. This is
also mass flow.
 TRANSLOCATION
EXPERIMENTS
1. Puncture expt.
If the phloem is punctured with a hollow tube then the
sap oozes out, showing that there is high pressure
(compression) inside the phloem (this is how maple
syrup is tapped). If the xylem is punctured then air is
sucked in, showing that there is low pressure (tension)
inside the xylem. This illustrates the main difference
between transport in xylem and phloem: Water is
pulled up in the xylem, sap is pushed down in the
phloem.
 2.Ringingexperiment
Since the phloem vessels are outside the xylem vessels,
they can be selectively removed by cutting a ring in a
stem just deep enough to cut the phloem but not the
xylem. After a week there is a swelling above the ring,
reduced growth below the ring and the leaves are
unaffected. This was early evidence that sugars were
transported downwards in the phloem.
 3.Aphidstyletexperiment
Aphids, such as greenfly, have specialised mouthparts
called stylets, which they use to penetrate phloem tubes
and sup of the sugary sap therein. If the aphids are
anaesthetised with carbon dioxide and cut off, the stylet
remains in the phloem so pure phloem sap can be
collected through the stylet for analysis. This surprising
technique is more accurate than a human with a syringe
and the aphids enzymes ensure that the stylet doesnt get
blocked.
 Summary
Materials translocated in phloem:
Translocated solutes are mainly carbohydrates
Sucrose is the most common translocated sugar
Phloem also contains:
Amino acids, proteins, inorganic ions, and plant hormones
Rate of translocation:
Movement in the phloem is rapid, well in excess of rates
of diffusion
Average velocity is 1 meter per hour
 Summary
Pathway of translocation:
Sugars and other organic materials are conducted
throughout the plant in the phloem by means of sieve
tube elements
Sieve tube elements display a variety of structural
adaptations that make the well suited for transport
Materials are translocated in the phloem from sources
(usually mature leaves) to sinks (roots, immature
leaves)
 Summary
The pressure-flow model of phloem translocation

At source end of pathway

Active transport of sugars into sieve cells
Water flows into sieve cells and turgor pressure increases

At sink end of pathway

Unloading (active transport again) of sugars
Water flows out of sieve cells and turgor pressure decreases

Flow is driven by a gradient of pressure.

Energy is required to establish the pressure gradient, but energy is not

required by cells of the pathway itself.

Animation