Fisiologi Tumbuhan

HUBUNGAN SOURCE DAN

SINK

Irfan Suliansyah
PS. Agroekoteknologi
Fakultas Pertanian Universitas Andalas
 TRANSLOCATION IN THE
PHLOEM

o What does it do?

o Pathways and conduits for translocation
sieve elements
- sieve tube elements (angiosperms)
- sieve cells (gymnosperms)
o Source-sink relationships
o The mechanism and water relations of
phloem translocation.
The phloem is the vascular system for moving (translocating)
sugars produced in photosynthesis (photosynthate) and other
substances throughout the plant.

0.5M
 Fig. 10.2

Bark
Phloem
Secondary phloem

Vascular cambium

Growth ring 3 (current year)

Xylem

Ring 2 (one year older)

Ring 3 (two years older)

Vascular bundle of clover

Fig. 10.1

Phloem

xylem
“Girdling” a woody plant causes swelling of stem
A classic
above theexperiment - girdling
point of damage, indicating a blockage
of phloem transport.
 More experimental evidence that phloem is the transport
tissue for carbohydrates.

Radioactive
labeling with
14
CO2 can trace
movement of
sugars in the
phloem, and from
source leaves
to sinks throughout
the plant.
Sampling the
phloem for
chemical analysis

Aphids insert a
feeding stylet into
phloem and this can
be used to collect
phloem exudate for
chemical analysis.
Sieve tube elements
• Tubular cells with end wall pores
and lateral sieve areas
• Membrane bound
• Have some organelles
• Have adjacent companion cells

Figure 10.3
Sieve element features
•living, membrane-bound cells (compare to tracheary
elements of xylem)

•lack some structures and organelles in most living

cells - no nuclei, vacuole, Golgi, ribosomes,
microtubules, microfilaments

•associated with companion cells that have full

set of structures and organelles

•have sieve areas or pores that interconnect

adjacent sieve elements
• angiosperm s.e. are called sieve tube elements, while
gymnosperms’ are called sieve cells (see Table 10.1 for differences).
Fig. 10.5

Cell wall between

sieve elements

Sieve plate pore

Companion cell
 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. Is passive or active.
Symplastic loading
Apoplastic loading
ATP-dependent sucrose
transport in sieve element
loading
Regulation of
sucrose loading?
 Phloem unloading occurs via
symplastic and apoplastic pathways
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
Loading and unloading of sugars
occurs in different veins
 How do metabolites move in the
phloem?

Pressure-driven bulk flow (Ernst Muench, 1930)

Source to sink

High phloem transport velocities (~ 100 cm h-1)

So diffusion is not involved

Pressure-flow model of translocation
in the phloem
 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.
PHLOEM UNLOADING via the apoplast and symplast.
4. 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. ΔΨS ↓ (more negative) → Water moves in → ΔΨP↑
2. At sink, unloading causes decrease in solute concentration, ΔΨS↑ (more
positive) → H2O leaves → ΔΨP↓
3. Solution moves by mass flow under pressure gradient from source to sink
In what direction does phloem transport
substances throughout the plant?

From an area of carbohydrate supply to an

area of carbohydrate demand.

Source ----> Sink

Source-sink relationships can explain the direction of
phloem translocation within the plant.
Source - produces more carbohydrates than required for its own needs
Sink - produces less carbohydrates than it requires
Anatomical and developmental determinants of the direction of source-sink
translocation.

1. Proximity - sinks tend to be supplied by closer sources

2. Vascular connections may cause distinct source-sink

patterns that counter proximity

3. Source-sink relationships may shift during development

Young leaf is
completely dependent
on carbohydrates from
other sources. It is
a strong sink.
As the leaf grows it
increasingly provides
for its own
carbohydrate needs.
Mature leaf is largely
a carbohydrate exporter (source)
Phloem transport
Velocities ≈ 1 m hour-1 , much faster than diffusion

What is the mechanism of phloem transport?

What causes flow?, What’s the source of energy?
 The pressure-flow model (Münch, 1930s)
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
 Sugars are moved from photosynthetic cells and
actively (energy) loaded into companion cells.

Fig. 10.14
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.

Fig. 10.14
 Phloem loading uses a
Fig. 10.16
proton/sucrose symport.
The pressure-flow model (Münch, 1930s)

Fig. 10.10
The pressure-flow model of phloem translocation
At source end of pathway
• Active transport of sugars into sieve cells
• s and w decrease
• Water flows into sieve cells and turgor increases

At sink end of pathway

• Unloading (active transport again) of sugars
• s and w increase
• Water flows out of sieve cells and turgor decreases
 Some key elements of phloem transport

Flow is driven by a gradient of pressure, P.

Energy is required to establish the pressure gradient,

but energy is not required by cells of the pathway itself.

Flow is in direction of higher total water potential,

(counter to direction water tends to flow passively).
 w
-1.1MPa

-0.4MPa