10/6/2014

Plants use the products of photosynthesis

in 2 ways

Fisiologi Tumbuhan Lanjut

BIO 4081

1.

new proteins, tissues, cells, structures growth & reproduction

2.

fuel for the above processes

ALOKASI DAN PARTISI BIOMASSA

In what direction does phloem transport substances

throughout the plant?
From an area of carbohydrate supply to an area of
carbohydrate demand.

Which way does photoassimilate flow?

Always follows the link from tissues producing it to

tissues that use it to power growth and development

Source Sink

Source: Daun dewasa dan jaringan yang aktif

berfotosintesis

Sink

: akar, jaringan batang, buah

source sink

Source actively photosynthesizing tissues

Net exporter: tissues can switch during development i.e.
leaves: young=sink, old=source

Sink storage tissues/growing tissues

Roots, young leaves, fruits

Net importer: not autotrophic

CHANGES IN SOURCE-SINK RELATIONS

DURING LEAF DEVELOPMENT
Translocation: movement of carbohydrates through vascular system

Mature Leaf

Young leaf

Partitioning : the relative amount of exported carbon used by

metabolic sinks within the plant

SINK

SOURCE

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Transport in plants

PATTERNS OF TRANSLOCATION: SOURCE TO SINK

How do plants transport carbohydrates?

Mass-flow or pressure-flow hypothesis:

1. Phloem sap is not translocated exclusively in either an upward

or downward direction and is not influenced by gravity; phloem
translocation occurs from source to sink.

Carbohydrates move from source (site produced or stored) to sink (site

used)

Carbohydrates actively transported into phloem at source

High concentration of carbohydrates causes greater osmotic
pressure in phloem; water moves in from adjacent xylem by
osmosis
Water influx creates (turgor) pressure inside phloem; pushing
water and dissolved carbohydrates through phloem
At sink, carbohydrates actively removed from phloem; reducing
osmotic pressure in phloem
water leaves phloem and reenters xylem, maintaining a
osmotic pressure gradient between sources and sinks

2. Sources include any exporting organ, typically mature leaves

exporting photosynthate.
3. Storage organs (roots, tubers, seeds, etc.) can also serve as
sources.
4. Sinks include any non-photosynthetic organ or tissue that does
not produce sufficient photosynthate to support its own
metabolic needs: roots, underground stems, buds, immature
leaves, flowers, fruits, etc.

The regulation of photoassimilate distribution into

plant organs governs their productivity
Important for yield of crop plants
Maximize yield of marketable part of crop plant
e.g., wheat farmers want more carbon to go to grain than
to roots
We can define two plant physiology terms that define how C is
allocated to plant parts
These terms are important in agronomy
They govern the destination of C in plants: allocation and
partitioning

Allocation

The regulation of the distributed of fixed carbon into

various metabolic pathways (i.e. the fate of fixed carbon)
include storage (starch), utilization (metabolic energy,
synthesis of other compound) and synthesis of transport
sugar (sucrose).
Allocation involves storage, metabolism and transport,
process which facilitate competition for photosynthates

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Allocation of fixed carbon is a balancing act between

providing energy and C skeletons
Synthesis of storage compounds. Starch is synthesized and
stored within chloroplasts and, in most species, is the primary
storage form that is mobilized for translocation during the
night. Plants that store carbon primarily as starch are called
starch storers.
Metabolic utilization. Fixed carbon can be utilized within
various compartments of the photosynthesizing cell to meet
the energy needs of the cell or to provide carbon skeletons for
the synthesis of other compounds required by the cell.

The plants competing processes for C include:

Leaf metabolism
Respiration of glucose provides:
ATP
Carbon skeletons for biosynthesis

Short-term storage
Plants can only photosynthesize in the light
But they need to grow at all times, therefore store carbon in
leaves (form is species dependent)

Synthesis of transport compounds. Fixed carbon can be

incorporated into transport sugars for export to various sink
tissues. A portion of the transport sugar can also be stored
temporarily in the vacuole

Translokasi di dalam sel parenkim

Starch: dicots
Sucrose: sugarcane, sugarbeet
Sucrose polymers (fructans): grasses, monocots

Leaf carbon storage is a buffer against environmental factors

affecting photosynthesis

A large amount of C allocation is destined for export

from the leaf to metabolic sinks

Immediate transport (export)

About of fixed carbon
Regulation of export poorly understood
Shifts during development
Sink (young leaves) little export
Source (mature leaves) switch off/down regulate
sucrose hydrolytic enzymes, increase SPS (sucrose
phosphate synthase)

For most plants:

Starch (made in chloroplasts): stored
Sucrose (made in cytoplasm): transported high synthesis
(SPS) activity, high export rate

Export of sucrose ~constant at night

Source leaves regulate allocation

Partitioning

Increase in photosynthesis rate in a source leaf results in

increased translocation rate from the source
Control point for allocation
Starch syntesis
Sucrose synthesis (including distribution of sucrose
between transport and temporary storage)
Regeneration of intermediates in the reduction cycle of
the Calvin Cycle

The differential distribution of photosynthates within the plant,

e.g. into different sinks; efficiency of partitioning from
vegetative sinks into storage organs translates into productivity
for many crops.
The differential distribution of photosyntates within plants
- Various sinks partition sugars
- Distribution must be balanced
- Many cultivars are economically inportant because the
partition to edible plant parts (fruits, grains)

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Exported carbon is partitioned between various sinks

Partitioning refers to the relative amount of exported carbon used
by metabolic sinks within the plant

Sinks compete for carbon!

Apical meristems
Young leaves
Fruit

Dependent on:
Vascular connections between source and sinks (best to have
direct links (vascular traces)
Same side of stem
As close as possible

How close sink is to source physically

Sink strength

Kemampuan relatif dari organ untuk menarik fotosintat

tergantung pada: ukuran & stadia pertumbuhan
Kedua faktor dan jarak menentukan laju aliran
fotosintat

Sink strength depends on sink size and activity

Sink strength = sink size x activity
sink size
= total biomass of the sink tissue
sink activity = rate of uptake of photosynthates per unit
biomass of sink tissue

Jarak dan laju menentukan gradien source: sink

Sink strength

Kemanpuan sink untuk memobilisasi fotosintat tergantung pada

ukuran dan aktivitas sink
Urutan kekuatan sink untuk hasil fotosintesis:
Biji > daging buah = pucuk daun muda > kambium > akar >
organ penyimpan lain

Source-sink pathways follow patterns

Proximity: of source to sink is a significant factor.
Upper nature leaves usually provide photosynthesis products to
growing shoot tip and young, immature leaves
Lower leaves supply predominantly the root system
Intermediate leaves export in both directions
Development: Importance of various sinks may shift during plant
development
Roots and shoots major sinks during vegetative growth
But fruits become dominant sinks during reproductive
development

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Source-sink pathways follow patterns

Vascular connections: Source leaves preferentially supply sinks
with direct vascular connections
A given leaf is connected via vascular system to leaves above and
below it on the stem

SOURCE TO SINK PATHWAYS FOLLOW ANATOMICAL AND

DEVELOPMENTAL PATTERNS
1. Proximity is important: upper mature leaves supply photosynthate to
growing shoot tip; lower leaves supply the root; middle leaves supply both.
2. Development influences transport:
a. Young leaves begin as sinks, gradually become sources.
b. Reproductive structures become dominant sinks during flowering.

Modifications of translocation pathways: - Interference with a

translocation pathway by mechanical wounding (or pruning)

vascular interconnections can provide alternate pathways for

3. Vascular connections important; source leaves preferentially supply sinks

to which they have vascular connections; typically, sources leaves supply
sinks along the same vertical row or orthostichy.

phloem transport
4. Phloem interconnections (anastomoses) can provide alternative pathways
in the event of wounding or pruning.

The sink strength of a plant organ is a measure of its

capacity to absorb photoassimilate
Sink strength = sink size (mass) X
sink activity (rate of uptake)
The sink strength of an organ defines
its capacity to assimilate
macromolecules
Why sink strength changes with time
or what determines it: unknown
Good:
High unloading rate
High sequestration rate into storage
Being close to source

Developing grain: very strong sink

No yield reduction even at low [CO2],
carbon stolen from roots and other
less strong sinks

Factors Affecting the Translocation

Temperature
Increased temperature increased loading & unloading optimum 20 30oC
Chilling Sensitive Plants (most)
Chilling Tolerant Plants (beets)
Can acclimate translocation of photosynthate to increasingly cold conditions

Factors Affecting the Translocation of Sap

Light
In the dark root translocation of photosynthate is favored
over stem translocation.
At least one study shows that the translocation of sap in
the stem was increased by BLUE and RED light.

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Factors Affecting the Translocation of Sap

Hormones
Both cell division (cytokinins) and cell elongation (auxins)
creates sinks absorbs sap.
Bud break
Increased GA, decreased ABA

Partitioning control is a complex process

The steps involved in C flow to a
sink: phloem loading in a
source leaf, phloem transport
into the sink, unloading and
short distance transport within
the sink and then metabolism
and storage in the sink.
- Molecule signalling: sucrose , hormone
- The status of various nutrient elements
affects the partitioning of photosynthate. n

The status of various nutrient elements affects the

partitioning of photosynthate
When plant growth is limited by a low supply of the
elements N, P, S or Fe, the ratio of root to shoot mass
rises, more organic C being delivered to the roots,
whereas limitation of the supply of Mg, Mn or K leads
to a decrease of the root : shoot mass ratio. Water
stress increases the root : shoot ratio, root growth
being stimulated while shoot growth is reduced.