Allocation, Translocation, and

Partitioning of Photoassimilates
 Learning objectives
• Understand the biochemical pathways that
lead to the synthesis of starch and sucrose.

• Know how carbon provided by photosynthesis is

allocated to starch or sucrose synthesis.

• Understand the nature of the phloem tissue and

its role in the transport of photoassimilates.
 Learning objectives
• Know the mechanism of phloem transport
and the importance of source-sink
relationships for transport.

• Recognize that some xenobiotic

compounds can enter the phloem and be
translocated internally.
 Starch and sucrose synthesis
• Photosynthesis produces simple hexose sugars
that are used by plants to synthesize starch and
sucrose.

• Whether those sugars are devoted to starch or

sucrose synthesis is called carbon allocation.

• Starch is a storage form of carbon while sucrose

is the primary form in which carbon is
transported over long distances in the plant.
 Terminology

• Carbon allocation: distribution of carbon

within the plant to different plant parts.
• Carbon partitioning: division of carbon
into metabolic, structural or storage pools
• Translocation is the movement of
materials from leaves to other tissues
throughout the plant.
 Starch synthesis
• Starch is synthesized in
the stroma of chloroplasts
in the leaves.

• There are two principle forms of starch.

– Amylose is a linear glucose polymer
constructed with a-(1,4) linkages.
– Amylopectin is similar to amylose except that it has a-
(1,6) linkages that create a branched chain structure.
 Starch synthesis
• The synthesis of
starch begins
with the hexose
phosphate pools
produced by the
PCR cycle.

(PCR =
Photosynthetic
Carbon Reduction)
 Starch synthesis
• Fructose-6-P is
converted to Glucose-
1-P by hexose-
phosphate isomerase
and
phosphoglucomutase.
• The Glucose-1-P
reacts with ATP to
form ADP-glucose,
mediated by ADP-
glucose
pyrophosphorylase.
 Sucrose and starch synthesis are
competing processes
• F-2,6-BP helps provide a balance between
CO2 assimilation and carbon allocation.
– Under low light conditions, sucrose export
decreases.
– Decreased export results in an accumulation
of F-6-P, triose phosphates, and F-2,6-BP.
– This accumulation decreases triose
phosphate export from the chloroplast.
– The increases in chloroplastic triose
phosphates and the decreased Pi stimulates
starch synthesis in the chloroplast.
 Sucrose synthesis
• The synthesis of the sucrose occurs in the
cytosol of photosynthetic cells by one of
two pathways.

• Unlike starch synthesis where glucose is

activated by ATP, glucose for sucrose
synthesis is activated by uridine
triphosphate (UTP) primarily, but also by
ATP.
 Sucrose synthesis
• Unlike starch synthesis where glucose is
activated by ATP, glucose for sucrose
synthesis is activated by uridine
triphosphate (UTP) primarily, but also by
ATP.
Sucrose synthesis
 Sucrose synthesis
• The principle pathway is mediated by sucrose
phosphate synthase and sucrose phosphate
phosphatase.

– UDP-glucose
reacts with
fructose-6-P to
form sucrose-6-P,
releasing UDP.
– Sucrose-6-P is
dephosphorylated
to form sucrose.
 Sucrose synthesis
• The carbon used to synthesize sucrose in the
cytosol is exported as DHAP from the
chloroplast on the Pi/triose phosphate
transporter.
 Sucrose synthesis

• G-3-P and DHAP

combine in the cytosol
to form fructose-1,6-
bisphosphate.
• The fructose-1,6-
bisphosphate enters
the cytosolic hexose
phosphate pool
where it is converted
to glucose-1-
phosphate.
 Sucrose and starch synthesis are
competing processes
• The principle factor determining the balance
between sucrose and starch synthesis is the
cytosolic enzyme fructose-1,6-
bisphosphate phosphatase (FBPase).

FBPase may
be an
ancestral Fructose 1,6-
gluconeogenic bisphosphate
enzyme
aldolase/phosphatase
Sucrose and starch synthesis are
competing processes
• This
enzyme
represents
a key
regulatory
point for
the flow of
carbon into
sucrose.

Figure 19-34
• Figure 19-34
• The concentration of the allosteric regulator fructose-
2,6-bisphosphate in plant cells is regulated by the
products of photosynthetic carbon fixation and by Pi.
• Dihydroxyacetone phosphate and 3 phosphoglycerate
produced by CO2 fixation inhibit phosphofructokinase-
2 (PFK-2), the enzyme that synthesizes the regulator;
• Pi stimulates that enzyme.
• The concentration of the regulator is therefore
inversely proportional to the rate of photosynthesis.
• In the dark, the concentration of fructose-2,6-
bisphosphate increases and stimulates the
glycolytic enzyme, PPi-dependent
phosphofructokinase-1 (PFK-1), while inhibiting
the gluconeogenic enzyme fructose-1,6-
bisphosphatase (FBPase-1).
• When photosynthesis is active (in the light), the
concentration of the regulator drops and the
synthesis of fructose-6-phosphate and sucrose is
favored.
• FBPase-2 represents fructose-2,6
bisphosphatase.
• The concentration of fructose-2,6-bisphosphate
varies inversely with the rate of photosynthesis
in higher plants.
• The enzyme phosphofructokinase-2, responsible
for fructose-2,6-bisphosphate synthesis, is
inhibited by dihydroxyacetone phosphate or 3-
phosphoglycerate and stimulated by Pi during
active photosynthesis, dihydroxyacetone
phosphate is produced and Pi is consumed,
resulting in inhibition of PFK-2 and lowered
concentrations of fructose2,6-bisphosphate.
 Sucrose and starch synthesis are
competing processes
• This type of photosynthetic control is said
to be feedback limited.
 Fructan biosynthesis
• Assimilated carbon can also be allocated to
fructan biosynthesis in the vacuole.
– Sucrose:sucrose fructosyl transferase uses to
sucrose molecules to form glucose and a
trisaccharide called 1-ketose.
– Fructan:fructan fructosyl transferase, a vacuolar
enzyme, extends the fructan polymer.

• Fructans can be synthesized when the rate of

carbon accumulation exceeds the rate of carbon
utilization.
 Fructan biosynthesis
• Fructan accumulators (chicory, wheat,
ryegrass, onion, agave, and pachysandra)
 Long distance transport of
photoassimilates
• Early experiments in which trees were
girdled provided some of the first
evidence that the phloem was the tissue
responsible for photoassimilate transport.

• Results from studies with aphids and

radiotracers (e.g., 14C) provided additional
information.
Long distance transport of
photoassimilates
Long distance transport of
photoassimilates
Composition of the phloem sap
• Phloem sap is a complex mixture of
organic and inorganic compounds.
– Sucrose and other sugars
– Proteins
– Amino acids
– Organic acids
– Anions and cations
– Phytohormones
Composition of the phloem sap
• Some plant species transport other sugars
in addition to sucrose.
– Some plants transport sugars from the
raffinose series.
– Some plants transport sugar alcohols such as
mannitol or sorbitol.

• Plants do not transport reducing sugars

in the phloem.
Composition of the phloem sap
Aphid acquiring
phloem sap
 Cellular constituents of the phloem
• The conducting cells of the
phloem are the sieve elements
(or sieve tubes).

• The individual cells, or sieve

tube members, are attached
end-to-end to form the phloem
network.

• The protoplast of these cells is

connected through sieve areas,
bordered between cells by the
sieve plate.
Cellular constituents of the phloem
• The sieve tube
members are highly
modified living cells.

• Associated with each

sieve tube member is
a companion cell.

• The companion cells

provide metabolic
support to the sieve
tube members.
Cellular constituents of the phloem
• Additional parenchymal cells associated
with the phloem are the transfer cells.

• These cells are likely involved in the

transfer of photoassimilates and other
compounds between mesophyll cells.
 Proteins in the phloem
• The phloem contains aggregates of proteins
called P-protein bodies.

• The role of P-proteins in phloem transport is

unclear but may involve plugging the sieve
plates of damaged cells.

• The glucan callose may serve a similar purpose

in sealing off damaged phloem cells.
Callose is also made in response of wounding
 Directionality of phloem transport
• Phloem transport moves
photoassimilates from
sources to sinks.
– A source is an area, such
as a storage organ or
photosynthetically active
leaf that is capable of
exporting
photoassimilates.
– A sink is an area that
must import
photoassimilates to
support metabolism.
Mechanism of phloem
transport
• The currently accepted model
for phloem transport is the
pressure flow hypothesis.

• According to this model,

phloem sap is driven by a
positive hydrostatic pressure
from source to sink.

• Phloem transport is tightly

linked with transpirational
water flow in the xylem.
Mechanism of phloem
transport
– Phloem loading occurs at
the source, as
photoassimilates are
transported into the sieve
tube members.
– The addition of these
solutes makes the water
potential of these cells
more negative.
– Water, following the water
potential gradient, diffuses
into the sieve tube
members from the xylem.
Mechanism of phloem transport
– The influx of water creates
a positive hydrostatic
pressure driving sap
movement.
– At the sink tissue, phloem
unloading occurs.
– This unloading makes the
water potential of the sieve
tube members less
negative.
– This change in water
potential causes water to
exit back to the xylem.
Mechanism of phloem transport
– The loading at the
source and unloading
at the sink creates a
differential that
maintains transport.
– Fundamentally, the
transport of the
phloem sap occurs
passively without the
direct input of energy.
Mechanism of phloem transport
• The driving force for phloem transport can
be simulated using osmometers and a
circuit of tubing.

• This system demonstrates that the

mechanism is not only feasible, but more
than adequate to provide the necessary
rates of transport.
Mechanism of phloem transport
• Since the phloem sap moves from source
to sink, and sources and sinks within a
plant change over time, then the phloem
must be bidirectional in its transport.

• This bidirectional transport may occur

through different vascular bundles, or even
through different sieve tubes within the
same bundle.
 Phloem loading and unloading
• Source cells (such as photosynthetic cells
in leaves) are generally located within a
few cells of a sieve element-companion
cell complex (se-cc).

• In leaves, sucrose is believed to move

through the symplasm via
plasmodesmata by diffusion to reach the
phloem parenchyma.
 Phloem loading and unloading
• There are two possible pathways from the
phloem parenchyma to the se-cc.
– A symplastic pathway may allow for transport directly
into the se-cc.
– Sucrose may also be transported into the cell wall
apoplasm and move via an apoplastic pathway to
the se-cc.
Phloem loading and unloading
 Phloem loading and unloading
• The transport of sucrose into the ss-cc
complex from the apoplasm would require
active transport.

• A sugar-H+ co-transport system, mediated

by genes such as SUT1 and SUC2, is
believed to be responsible for this
transport step.
Phloem loading and unloading
Phloem loading and unloading
 Phloem loading and unloading
• Sucrose moving symplastically into the se-cc is
converted to oligosaccharides.

• The synthesis of these oligosaccharides

prevents back-diffusion of sucrose into the
source cells and maintains the concentration
gradient into the se-cc.
 Phloem loading and unloading
• Phloem unloading is, in principle, the same as
phloem loading except for the directionality.

• Both symplastic and apoplastic pathways are

believed to occur.

• The symplastic pathway has been observed in

young, developing leaves and root tips, and
involves sucrose diffusion followed by hydrolysis
of the sucrose.
 Phloem loading and unloading
• There are two possible apoplastic
pathways for phloem unloading.
– Sucrose released into the apoplast can be
hydrolyzed into glucose and fructose by acid
invertase, and the monosaccharides are then
taken up by sink cells.
– Sucrose can be transported into the apoplasm
by an energy-dependent carrier.
 Link animasi phloem loading
• https://highered.mheducation.com/sites/98
34092339/student_view0/chapter38/anima
tion_-_phloem_loading.html
 Allocation and partitioning
• While some of the photoassimilates are
retained within the photosynthetic leaves,
the rest can be exported.

• The distribution of those exported

photoassimilates to various processes
involves allocation and partitioning.
 Allocation and partitioning
• Allocation is the metabolic fate of
photoassimilates.

• Photoassimilates may be retained in the

photosynthesizing leaf to support metabolism.

• Some of the photoassimilates are stored in the

photosynthesizing leaves.

• The bulk of the photoassimilates are exported.

 Allocation and partitioning
• The distribution of photoassimilates
between sinks is called partitioning.

• The partitioning is determined by

competition between sink tissues.

• The sink strength determines which sink

obtains a larger fraction of the
photoassimilate pool.
 Allocation and partitioning
• Sink strength is a function of sink size and
sink activity.
– Sink size refers to the biomass of the sink
tissue (usually as dry weight).
– Sink activity is the rate of uptake of assimilate
per unit biomass per unit time.
 Allocation and partitioning
• Sink strength is also a function of:
– Proximity of the sink to the source
– Environmental factors
– Cell turgor
– Hormones
• Source organs that are usually
photosynthetically active are defined as
net exporters of photoassimilates,
represented mainly by mature leaves,
• Sink organs that are photosynthetically inactive are
referred to as net importers of fixed carbon.
• Sinks divided into two different classes: utilization and
storage sinks.
• Utilization sinks are highly metabolically active, rapidly
growing tissues such as meristems and immature leaves
• Storage sinks are the organs like tubers, seeds and
roots, where the imported carbohydrates are deposited
in the form of storage compounds (e.g. starch, sucrose,
fatty acids, or proteins)
• The storage sinks are usually specialized
for other essential processes, such as
mineral acquisition (roots) or reproduction
(seeds, fruits and potato tubers).
• Metabolic sink or source status of a
particular organ is under developmental
control.
 Xenobiotics in the phloem
• Synthetic chemicals, or xenobiotics, with
the appropriate lipophilicity, can also be
transported in the phloem.

• The agrochemical industry has an interest

in understanding this process because it
relates to the internal distribution of
herbicides, pesticides, and growth
regulators.
 Xenobiotics in the phloem
• One example of a phloem-mobile xenobiotic is
the herbicide glyphosate.
• Glyphosate applied to leaves enters the
phloem and is transported to roots where it
interferes with aromatic amino acid
metabolism.
 Glyphosate
• Glyphosate is a non-selective herbicide,
meaning it will kill most plants.
• It prevents the plants from making certain
proteins that are needed for plant growth.
• Glyphosate stops a specific enzyme
pathway, the shikimic acid pathway.
• Plants absorb glyphosate through their
leaves and other green parts. From here,
the glyphosate moves to the growing
points of shoots and roots, where it
interferes with the enzymatic production of
certain amino acids that are essential for
plant growth.
• This pathway exists only in plants, fungi
and bacteria, so the toxicity to animals is
low.