ADP + Pi

ATP
Symplast

H+ Sucrose

Cell membrane

Apoplast
H+

H+ Sucrose

9
Allocation, Translocation, and Partitioning
of Photoassimilates
The previous two chapters showed how energy was
conserved in the form of carbon compounds, or photoassimilates. The primary function of photosynthesis
is to provide energy and carbon sufcient to support
maintenance and growth not only of the photosynthetic
tissues but of the plant as a whole. During daylight
hours, photoassimilate generated by the PCR cycle is
temporarily accumulated in the leaf as either sucrose
in the mesophyll vacuole or starch in the chloroplast
stroma. The conversion of photoassimilates to either
sucrose or starch is called carbon allocation. Although
a portion of the carbon assimilated on a daily basis is
retained by the leaf to support its continued growth
and metabolism, the majority is exported out of the
leaf to nonphotosynthetic organs and tissues. There, it
is either metabolized directly or placed in storage for
retrieval and metabolism at a later time. The transport of photoassimilates over long distances is known
as translocation. Translocation occurs in the vascular tissue called phloem. Phloem translocation is a
highly signicant process that functions to ensure an
efcient distribution of photosynthetic energy and carbon between organs throughout the organism. This
is called carbon partitioning. Phloem translocation is
also important from an agricultural perspective because
it plays a signicant role in determining productivity,

crop yield, and the effectiveness of applied herbicides

and other xenobiotic chemicals.
This chapter is focused on the biosynthesis of
the photosynthetic end-products, starch and sucrose,
and the structure of the phloem and its function in
the translocation and distribution of these important
photoassimilates. The principal topics to be covered
include

151

the biosynthesis of starch and sucrose,

the allocation of xed carbon between the starch
and sucrose biosynthetic pathways,
the basis for identifying phloem as the route for
translocation of photoassimilates and the nature of
substances translocated in the phloem,
the structure of the phloem tissue, especially the
several unique aspects of sieve tube structure and
composition,
the source-sink concept and the signicance of
sources and sinks to the translocation process,
the pressure-ow hypothesis for phloem translocation and the processes by which photoassimilates
gain entry into the phloem in the leaf and are subsequently removed from the translocation stream at
the target organ,

152

Chapter 9 / Allocation, Translocation, and Partitioning of Photoassimilates

factors that regulate the distribution of photoassimilates between competing sinks, and
the loading and translocation of xenobiotic agrochemicals.

9.1 STARCH AND SUCROSE

ARE BIOSYNTHESIZED
IN TWO DIFFERENT
COMPARTMENTS
Many plants, such as soybean, spinach, and tobacco,
store excess photoassimilate as starch in the chloroplast,
while others, such as wheat, barley, and oats, accumulate little starch but temporarily hold large amounts of
sucrose in the vacuole. The appropriation of carbon
xed by the PCR cycle into either starch or sucrose
biosynthesis is called carbon allocation. This will be
discussed further in relation to source sink relationships (see below). The starch and sucrose will later be
mobilized to support respiration and other metabolic
needs at night or during periods of limited photosynthetic output. Sucrose exported from the leaf cell to
nonphotosynthetic tissues may be metabolized immediately, stored temporarily as sucrose in the vacuoles,
or converted to starch for longer-term storage in the
chloroplasts.

9.1.1 STARCH IS BIOSYNTHESIZED

IN THE STROMA
The dominant storage carbohydrate in higher plants
is the polysaccharide starch, which exists in two forms
(Figure 9.1). Amylose is a linear polymer of glucose
created by linking adjacent glucose residues between
the rst and fourth carbons. Amylose is consequently
known as an -(1,4)-glucan. Amylopectin is similar to
amylose except that occasional -(1,6) linkages, about
every 24 to 30 glucose residues, create a branched
molecule. Amylopectin is very similar to glycogen, the
principal storage carbohydrate in animals. Glycogen is
more highly branched, with one -(1,6) linkage for
every 10 glucose residues compared with one in 30 for
amylopectin.
The site of starch synthesis in leaves is the
chloroplast. Large deposits of starch are clearly evident in electron micrographs of chloroplasts from
C3 plants. In addition, the two principal enzymes involvedADPglucose pyrophosphorylase and starch
synthaseare found localized in the chloroplast
stroma. Starch synthesis in the chloroplast begins with
the hexose phosphate pool generated by the PCR
cycle (Figure 9.2). Fructose-6-phosphate (F-6-P), one
component of the stromal hexose phosphate pool, is
converted to glucose-1-phosphate, another component
of the stromal hexose phosphate pool, by the two

CH2OH
CH2OH
O H
O H
H
H
H
OH H
OH H
O
O
H

OH

Glucose

OH

Glucose
n

-Amylose

Branch

...

CH2OH
CH2OH
O H
O H
H
H
H
OH H
OH H
O
O
H OH
H OH

(1 6) Branch point

CH2
CH2OH
CH2OH
CH2OH
O H
O H
O H
O H
Main H
H
H
H
H
H
H
H
chain
OH H
OH H
OH H
OH H
... O
O
O
O
O
H

OH

OH

OH

OH

Amylopectin

FIGURE 9.1 The chemical structures of the two forms of starch: amylose and amylopectin. Amylose is a long chain of (14)-linked glucose residues. Amylopectin
is a multibranched polymer of (14)-linked glucose containing (16) branch
points.

...

153

9.1 Starch and Sucrose Are Biosynthesized in Two Different Compartments

Light

CO2
MESOPHYLL CELL

H
Cytosol

CO2

HO

Pi

HO

Pi

Triose-P

CH2OH H

Triose-P
Translocator

Triose-P

Hexose-P
pool

OH
O
HO
Sucrose

Pi
Hexose-P
pool

H
CH2OH
H
O HO

B.

CH2OH

Pi

Pi

Starch

Sucrose

Translocation

Chloroplast

A.

FIGURE 9.2 (A) Allocation of xed carbon between the chloroplast and the cytosol.
(B) The structure of sucrose.

chloroplast enzymes, hexose-phosphate isomerase

(Equation 9.1) and phosphoglucomutase (Equation 9.2).
The glucose-1-P subsequently reacts with ATP to form
ADP-glucose (Equation 9.3). Reaction 9.3 is catalyzed by the enzyme ADP-glucose phosphorylase.
ADP-glucose is an activated form of glucose and serves
as the immediate precursor for starch synthesis. Starch
deposits within the chloroplast stroma are evident as
insoluble starch grains. As a consequence, this form of
stored carbon is osmotically inactive (Chapter 1) which
allows plants to store large amounts of xed carbon
in chloroplasts with minimal inuence on the osmotic
pressure of the stroma. This prevents the chloroplast
membrane from bursting upon the accumulation and
storage of xed carbon as starch.
fructose-6-P glucose-6-P

(9.1)

glucose-6-P glucose-1-P

(9.2)

ATP + glucose-1-P ADP-glucose + H2 O + PPi

(9.3)
PPi + H2 O 2Pi

(9.4)

Finally, the enzyme starch synthase catalyzes formation

of a new -(1,4) link, adding one more glucose to the
elongating chain (Equation 9.5).
ADP-glucose + -(1 4)-glucan ADP
+ -(1 4)-glucosyl-glucan (9.5)
Formation of the -(1,6) branching linkages, giving rise
to amylopectin, is catalyzed by the branching enzyme,
also known as the Q-enzyme.

9.1.2 SUCROSE IS BIOSYNTHESIZED

IN THE CYTOSOL
Sucrose is a soluble disaccharide containing a glucose
and a fructose residue (Figure 9.2B). It is one of the more
abundant natural products that not only plays a vital role
in plant life but is also a leading commercial commodity.
Sucrose may function as a storage product as it does in
sugarbeets or sugarcane, where it is stored in the vacuoles
of specialized storage cells. Alternatively, sucrose may
be translocated to other, nonphotosynthetic tissues in
the plant for direct metabolic use or for conversion to
starch. Sucrose is by far the most common form of sugar
found in the translocation stream.
The site of sucrose synthesis in the cell was the
subject of debate for some time. On the basis of
cell fractionation and enzyme localization studies it
has now been clearly established that sucrose synthesis occurs exclusively in the cytosol of photosynthetic
cells (Figure 9.2). Earlier reports of sucrose synthesis
in isolated chloroplasts appear attributable to contamination of the chloroplast preparation with cytosolic
enzymes. Moreover, the inner membrane of the chloroplast envelope is impermeable to sucrose, so that if
sucrose were synthesized inside the chloroplast it would
be unable to exit the chloroplast and enter the translocation stream.
Two routes of sucrose synthesis are possible. The
principal pathway for sucrose synthesis in photosynthetic cells is provided by the enzymes sucrose phosphate synthase (Equation 9.6) and sucrose phosphate
phosphatase (Equation 9.7).