Translocation in the Phloem

Phloem and xylem are structures for long-distance

transport

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 Phloem is always in close proximity to xylem.

Root Stele Stem Vascular Bundle Leaf Midrib

Primary phloem and primary xylem

Stem of
Clover

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 Secondary phloem and secondary xylem
3-year-old stem of
an ash tree

(Growth rings)

Classical studies on phloem transport

Marcello Malpighi in 1686:

- removed bark of a tree in a
ring around the trunk
(girdling);
- no immediate effect on
transpiration
- sugar transport in the trunk
is blocked at the site where
the bark has been removed
- Sugars accumulate above
the girdle (on side toward the
leaves) and are depleted
below the treated region

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 Sugar is translocated in the phloem
Phloem consists of 3 types of cells:
- Sieve element (sieve tube element,
sieve cell)
- Companion cell
- Phloem parenchyma

Only sieve elements are

involved in translocation.

Sugar is translocated in phloem sieve elements

Sieve elements: conduct sugars and other organic materials

throughout the plant

Sieve elements

Sieve tube elements Sieve cells

in angiosperms in gymnosperms

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 Schematic drawings of mature sieve elements (sieve tube elements)

A rostacsövet alkotó sejtekben

megtalálható sejtalkotók:
módosult MT és PL, SER
hiányzó sejtalkotók: SM,
mikrofilamentumok,
mikrotubulusok, vakuólum,
Golgi, riboszómák

A sejtfalra jellemző a
másodlagos vastagodás,
ami viszont nem lignifikált

A rostacsõtagok végfalai
az ú.n. rostalemez:
pórusok átmérõje 1-15 µm

Sieve tube elements and companion cells

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 Sieve areas of angiosperms can differentiate into
sieve plates

Sieve elements in
winter squash

Sieve area linking two sieve cells of a pine

sa = sieve area
P = plastid
SER = smooth ER

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 Three different types of companion cells
Ordinary companion cells
- have chloroplasts
- few plasmodesmata between companion cell and
surrounding cells, except for own sieve elements
- symplast of sieve element and its companion cell is relatively
isolated from surrounding cells
Transfer cells
- similar to ordinary companion cells
- develop fingerlike wall ingrowths, particularly on walls that face
away from sieve element
- wall ingrowths increase surface area of plasma membrane
(increases potential for solute transfer across membrane)
Intermediary cells
- have numerous plasmodesmata connecting them to bundle sheath cells
- have many small vacuoles
- poorly developed thylakoids and lack of starch grains

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Three different types of companion cells
Scarlet monkey flower

Three different types of companion cells

Pea

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 Three different types of companion cells
Maskflower
Plasmodesmata

Patterns of translocation: Source to Sink

Metabolites move from source to sink

SOURCE = area of supply

- exporting organs: mature leaves
- storage organs: seed endosperm, storage
root of second growing season beet

SINK = areas of metabolism (or storage)

- non-photosynthetic organs and organs that
do not produce enough photosynthetic
products to support their own growth or
storage
- Example: roots, tubers, developing
fruits/seeds, immature leaves

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Source-to-sink patterns of phloem translocation

A B
Source leaf

Sugar Beet

Source-to-sink patterns of phloem translocation

source leaf

Sugar Beet

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 Transition form sink to source is gradual

A-D: leaf imported 14C from

the source leaf on the plant
for 2h

Label is visible as black

accumulation

Translocation of Substances in the Phloem

• Translocation (movement of organic solutes) stops if the

phloem is killed.
• Translocation often proceeds in both directions— both up
and down the stem simultaneously.
• Translocation is inhibited by compounds that inhibit
respiration and the production of ATP.
• Velocities ≈ 1 m hour-1 , much faster than diffusion

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Aphids = Phloem miners

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 Translocation of Substances in the Phloem

• Plant physiologists have used aphids to collect sieve tube

sap from individual sieve tube elements.
• An aphids inserts a specialized feeding tube, or stylet,
into the stem until it reaches a sieve tube.
• Sieve tube sap flows into the aphid. The aphid is then
frozen and cut away from its stylet, which remains in the
sieve tube.
• Sap continues to flow out the sieve tube and can be
collected and analyzed by the physiologist.

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Structures of compounds are not translocated in the phloem

Structures of compounds commonly translocated in the phloem

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Structures of compounds commonly translocated in the phloem

Phloem loading and unloading

Source:
PHLOEM LOADING is energy-dependent transport.
Sucrose is actively loaded into companion cell or phloem
parenchyma cells by H+-coupled symport.
Symplast or apoplast; can also be passive.

Sink:
PHLOEM UNLOADING is via the apoplast and symplast. It is
passive or active.

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 Summary apoplastic and symplastic loading
• Species with apoplastic loading
– Mainly sucrose transport
– Herbs, temperate or arid climate plants
• Species with symplastic loading
– Mostly intermediary companion cells
– Transport raffinose and stachyose
– Trees, shrubs, tropical and subtropical plants
• Species with both types of loading
– Coleus

Sugars are moved from photosynthetic cells and actively

(energy) loaded into companion cells.

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Sugars are moved from photosynthetic cells and actively
(energy) loaded into companion & sieve cells.
The concentrating of sugars in sieve cells drives the osmotic
uptake of water.

Apoplastic loading

Symplastic loading

ATP-dependent sucrose
transport in sieve element
loading

Phloem loading uses a

proton/sucrose symport

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Phloem unloading occurs via symplastic and
apoplastic pathways

Loading and unloading of sugars occurs in

different veins

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 Physiological
process of loading
sucrose into the
phloem

Pressure-flow
Phloem and xylem are coupled
in an osmotic system that
transports sucrose and
circulates water.

Physiological process
of unloading sucrose
from the phloem into
the sink

Translocation of Substances in the Phloem

• Sieve tube cells at the source have a greater sucrose

concentration that surrounding cells, so water enters by
osmosis. This causes greater pressure potential at the
source, so that the sap moves by bulk flow towards the
sink.
• At the sink, sucrose is unloaded by active transport,
maintaining the solute and water potential gradients.
• This is called the pressure flow model (Münch, 1930s).

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Pressure-flow model of translocation in the phloem

Water Cycling

Pressure-flow hypothesis
Source
• High concentration of sucrose, via
photosynthesis,
– ∆[sucrose] drives diffusion,
• Active H+-ATPase,
– electrochemical gradient drives
symporters,
• - Ψs builds, water enters the cell
• + Ψp builds.

Sink
• Low concentration of sucrose,
– ∆[sucrose] drives diffusion,
• Active H+-ATPase,
– electrochemical gradient drives
antiporters,
• - Ψs drops, water exits the cell
• + Ψp drops.

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 Pressure-flow hypothesis

Ψp

• Notice that the Ψs at the source is

more negative than at the sink
• Why don’t we expect water to flow
toward the source?

Water, along with solutes moves

down the pressure gradient, not the
water potential gradient.

Allocation and partitioning

Allocation = the regulation of the distribution of fixed carbon

into various metabolic pathways (i.e. the fate of
fixed carbon)
- includes storage (starch), utilization (metabolic
energy, synthesis of other compounds), and
synthesis of transport sugars
Partitioning = the differential distribution of photosynthates
within the plant
- various sinks partition sugars
- distribution must be balanced
- Many cultivars are economically important because
they partition to edible plant parts (fruits, grains)

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 Source leaves regulate allocation

Increase in photosynthesis rate in a source leaf results in

increased translocation rate from the source.

Control points for allocation:

- starch synthesis
- sucrose synthesis (including distribution of
sucrose between transport and temporary
storage)
- regeneration of intermediates in the reduction
cycle of the Calvin Cycle

Source leaves regulate allocation

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Sink tissues compete for available translocated
photosynthates

Example: young leaves compete with roots for

photosynthates
In sugar beet and bean:
– Over short term: rates of photosynthesis and export from
single source do not change; young leaves receive
relatively more sugar than roots
– Shading decreases partitioning in roots
– Young leaves can deplete sugar content of sieve
elements more rapidly, thus increasing the pressure
gradient and translocation toward themselves

Sink Strength depends on sink size and activity

Sink strength = sink size × sink activity

» Sink size = total biomass of the sink tissue
» Sink activity = rate of uptake of photosynthates
per unit biomass of sink tissue
Plant types and storage strategies
Annuals, plants that usually germinate, flower and die
in one year (pea)
Biennials, plants that take between twelve and twenty-
four months to complete their lifecycle (carrot, celery,
parsley)
Perennials, plants that live for more than two years
(trees, shrubs)

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 Environmental influences on partitioning

• Water stress
– More root growth
• Nitrogen
– High N supply…less root growth
– Low N supply…more root growth

Transport of signal molecules

• Growth regulators (plant hormones, e.g. abscisic acid,

gibberellins, cytokinin)
– Affect partitioning by controlling sink growth, leaf
senescence etc.
– Abscisic acid enhances, auxin inhibits sucrose uptake by
some sink tissue
• Proteins (P-proteins, H+/sucrose symporter)
• Turgor pressure may control activities of sources and sinks
– e.g. phloem unloading is rapid due to rapid sugar
utilization at sink, ΨP ↓ in sieve elements; this reduction in
pressure is transmitted to source, i.e. loading increases

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 Phloem transport summary

1. Metabolites are transported from source to sink.

SOURCE: Tissue/organ that makes or stores food reserves. A net exporter.
E.g. Seed endosperm, Green Photosynthetic leaf.
SINK: Organ/tissue or cell that requires metabolites for energy and for
biosynthesis. A net importer. E.g. Shoot meristem, roots, developing seeds.
2. Phloem contains 3 types of cells.
3. PHLOEM LOADING in source tissue is by energy-dependent transport.
Sucrose is actively loaded into companion cell or phloem
parenchyma
by H+- coupled symport.
4. PHLOEM UNLOADING via the apoplast and symplast.

Phloem transport summary

5. LONG DISTANCE MOVEMENT in phloem is driven by a pressure

gradient.
1) At source, accumulation of sugars in
sieve elements → high solute concentration,
i.e. ∆ΨS ↓ (more negative) → water moves
in → ∆ΨP↑
2) At sink, unloading causes decrease in
solute concentration, ∆ΨS↑ (more positive)
→ H2O leaves → ∆ΨP↓
3) Solution moves by mass flow under
pressure gradient from source to sink.

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