PHLOEM TRANSPORT OF ASSIMILATES

AND ITS REGULATION

INTRODUCTION
• Non-photosynthetic tissues and organs of a plant need to be supplied
with energy and fixed carbon. During the process of photosynthesis
carbon is fixed and is stored in the form of starch and sugars
(photoassimilates). Starch is biosynthesized in the stroma region while
sugars in the cytosol. Sugars are also produced during the breakdown
of starch. Sugars, synthesized in the mesophyll cells, serve as the major
exported photosynthetic product. The photoassimilates are then
translocated to other non-photosynthetic tissues for metabolism or for
storage. Efficient transport of sugars across plants organs is a major
factor affecting plant growth

• Source-sink relationship:
During the process of photosynthesis or during the breakdown of starch,
sugar is produced in plants. A plant organ that produces sugar is termed as
source. While a plant organ in which sugar is either deposited or consumed,
is termed as sink. Sugar flows from source to sink and always follows a
shortest path, that is, it flows from nearest source to sink. This movement of
sugar from source to sink is termed as translocation. The process of
translocation takes place through phloem. It is believed that phloem sap
moves through sieve tubes by bulk flow rather than by diffusion, since
diffusion is inefficient for transport over long distances. Hence bulk flow may
have evolved as an efficient method of long distance transport in plants. The
sap moves at the rate of about 1 meter/hour.
PHLOEM TRANSPORT OF ASSIMILATES:
• Phloem loading plays a central role in determining productivity. While
most photoassimilate loading occurs in photosynthetically active
leaves, root-produced metabolites, such as amino acids, move readily
from xylem to phlom particularly at the stem nodes. Pholem tissue:
Phloem is a complex tissue composed of various specialized cells called
sieve tubes, companion cells, phloem fibres and phloem parenchyma

• 1. Sieve tubes - Sieve tube elements are elongated

schlerenchyma cells. A series of sieve tube elements are joined
end to end to form a tube like structure. The cross wall between
two adjacent elements become perforated to form sieve plates.
The cell walls are thin and the lumen is filled with slimy sap
composed of mainly proteins.

• 2. Companion cells- These are specialized parenchyma cells

which are elongated and thin walled with a distinct nucleus.
They are connected with sieve tubes by small canal.

• 3. Phloem fibres- These are elongated tapering cells which

have thick walls. These cells are found in stem.

• 4. Phloem parenchyma- The cells have thin walls and form a

packing tissue between all other types of cell

• Pressure flow hypothesis

• : Pressure flow hypothesis or Mass flow hypothesis has been
experimentally supported to explain movement of sugars through
phloem.The pressure-flow model was first proposed byErnst Münch in
1930. A turgor pressure gradient is created due to building of pressure
at the source end and gradual decrease in pressure towards sink end.
The movement of water and solutes takes place from source to sink
because of this pressure gradient. This pressure flow hypothesis may
not be true in non-flowering vascular plants, since they have small or
blocked phloem cell pores.
 Step 1 : At the source, sugar is loaded (active transport) into the
companion cell of phloem in leaf veins.It raises concentration of sugars in
companion cells above that in the sieve tube elements.

Step 2: Now sugar molecules move into sieve tube elements by diffusion
leading to an increased concentration of sugars in sie ve tube element.As a
result, there is decrease in water potential which leads to water uptake by
osmosis.

Step 3: A positive ‘fluid pressure’ is created due to water uptake than the
pressure at distant locations (near the sink).This pressure difference causes
movement of phloem sap down through sieve tube.

Step 4: Sugar is unloaded at the sink leading to decrease in pressure at that

location. Thus the procedure to produce pressure gradient continues leading
to bulk flow of sugars to the sink.

• In angiosperms sugar transport is facilitated through sieve tubes. It

takes place by active transport because sugar concentration in phloem
is high as compared to surrounding mesophyll cells. Simultaneous with
the translocation of sugar from source (leaves) to sink (roots),
movement of water also takes place from sink to source.

• Regulation of phloem transport:

• Transport of photo assimilates through phloem is an important
interface between photo assimilate production by leaves (sink) and
photo assimilate use by importing sink regions. Rate of phloem
transport is regulated by both source and sink processes. Though
transport through sieve tubes takes place from source to sink, the
pathway is affected by various factors. Phloem transport depends on
proximity, development, vascular connections, and modification of
translocation pathways. Also certain sources prefer to supply specific
sinks. The importance of various sinks may shift during plant
development. Shoot and root apices are usually the major sinks during
 vegetative growth, while seeds and fruits are the dominant sinks
during reproductive development, particularly for adjacent or nearby
leaves. In a source-limited system, does not elicit immediate effect on
phloem loading as it depends upon the capacity of leaves to increase
the size of transport pool. In contrast, sink-limited system, elicits
immediate effects on photo assimilate export.

• A. Internal factors \
• 1. Proximity of the source: The proximity of the source to the
sink is a significant factor. The upper mature leaves on a plant usually
provide photo assimilates to the growing shoot tip and young,
immature leaves; the lower leaves supply predominantly the root
system. Intermediate leaves export in both directions, bypassing the
intervening mature leaves. Source leaves preferentially supply sinks
with which they have direct vascular connections.

• 2. Regulation by the source/ sink:

• Transport of photo-assimilates depends on source supply and sink
demand. With increase in the net photosynthetic rate, export of sugar
(triose phosphate) from chloroplasts also increases, leading to feed-
forward up-regulation of sucrose biosynthesis. As a result, there is
amplification of sucrose transport pool. Activity of sucrose phosphate
synthetase (SPS) enzyme regulates the size of sucrose transport pool.
Activity of SPS is in turn, under the control of sink and environmental
factors. (Plants in action, chapter 5)When net photosynthetic rate is
low, SPS enzyme activity will not be inhibited by feedback inhibition by
substrates. As a consequence, there is an increase in photosynthetic
enzyme activity leading to increase in phloem transport to fulfill sink
demand. But high sucrose contents in mesophyll cells may have an
inhibitory effect on SUT activity and phloem loading and in turn down
regulation of photosynthesis. Sink strength or sink dominance is the
ability of a sink to effectively lower its sugar concentration so that a
 favourable hydrostatic pressre gradient between source and sink is
maintained

• 3. Growth hormones:
• Phytohormone levels in leaves respond to changes in the source/sink
ratio. Changes in the turgor pressure of phloem sap or altered
phytohormone levels could serve as signals for sink demand.
Hormones such as auxin and cytokinin have long been known to
increase the rate of phloem transport. Fusicoccin and auxin can rapidly
promote sucrose uptake, whereas abscisic acid acts as an inhibitor. (La
londe et al, 1999) Levels of gibberellin in the leaves near a developing
flower or a fruit increase. When auxin or Gibberellic acid was directly
applied to the leaves, it was observed that there was increase in
photoassimilate export rate but not in the rate of photosynthesis. So it
appears that gibberellic acid up-regulates phloem loading.

• B. Environmental factors:
•

• Sugar transport through the phloem can be affected by many

environmental factors that alter source/sink relationships. Several
abiotic factors that affect phloem transport include water and salt
stress, mineral deficiency, CO2, light, temperature, air, and soil
pollutants, while biotic factors include mutualistic and pathogenic
microbes, viruses, aphids and parasitic plants.

• Abiotic factors
•

• Water and salt stress: Drought imposes unfavorable conditions

on the leaves (source) and roots (sink) of a plant. However, as pointed
out by Turgeon (2010), the high osmotic potential in the phloem can be
 a positive parameter for attracting water to the sieve tubes and
maintaining phloem sap flow in drought conditions. Osmotic stress
promoted sucrose biosynthesis instead of starch biosynthesis via the
induction of sucrose-phosphate synthase (SPS).Sugar accumulation
may have a role in osmotic adjustment. Effect of water deficit is acute
in the reproductive stage as sugars allocation to flowers, seeds or f
ruits, is compromised by its allocation to roots.

Mineral deficiency: In case of shortage of minerals, plants allocate more

resources to organs involved in mineral acquisition. So nutrient deficiency
can affect photo-assimilate partitioning either directly via phloem loading
and transportor in directly by depressing sink demand. Nitrogen deficiency
reduces photosynthesis by a decrease in RubisCO amount and activity and
also a decrease in electron transfer. Lack of phosphorus in leaf mesophyll
cells has a direct effect on photosynthesis through Pi availability in the
chloroplast and leads to reduced carbon assimilation .Enhance
dcarbohydrate transport to the roots has been demonstrated for N and P
limitation, but not for K or Mg deficiency and the underlying mechanisms
have been partly unraveled. Polyols are considered as major molecules for
plants to cope with stress. They act both as osmotically active and anti-
oxydant molecules.Long- distance polyol transport enhances in response to
salt stress from shoot to root. Delivery of polyols to roots could have a
positive effect on metabolism and water potential of roots.