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