Sucrose Synthesis

Vijendra Shekhawat
UDP-Glucose is the Substrate for Sucrose Synthesisin the Cytosol of Leaf Cells
Most of the triose phosphate generated by CO2 fixation in plants is converted to
sucrose or starch. Sucrose may have been selected as the transport form of carbon
because of its unusual linkage between the C-1 of glucose and the C-2 of fructose. This
bond is not hydrolyzed by amylases or other common carbohydrate-cleaving enzymes,
and the unavailability of the anomeric carbons prevents sucrose from reacting non-
enzymatically (as does glucose) with amino acids and proteins.

Sucrose is synthesized in the cytosol, beginning with dihydroxyacetone phosphate and
glyceraldehyde 3-phosphate exported from the chloroplast. After condensation of two
triose phosphates to form fructose 1,6-bisphosphate (catalyzed by aldolase), hydrolysis
by fructose 1,6-bisphosphatase yields fructose 6-phosphate.

Sucrose 6-phosphate synthase then catalyzes the reaction of fructose 6-phosphate with
UDP-glucose to form sucrose 6-phosphate. Finally, sucrose 6-phosphate phosphatase
removes the phosphate group, making sucrose available for export to other tissues.

Chloroplast- Calvin Cycle- Triose Phosphate


Degradation Of Sucrose: Both invertase and sucrose synthase can breakdown sucrose:
Plant tissues contain distinct invertases located in the vacuole, cell wall (acid invertases) cytosol,
mitochondria, nucleus, and cholorplast (neutral/alkaline invertases) which hydrolyse sucrose to glucose
and fructose in an irreversible reaction.
Multiple isoforms of sucrose synthase are located in the cytosol or cytosolic membranes that catalyse a
thermodynamically reversible reaction, this reaction probably acts only to breakdown sucrose in vivo.



Synthesis and Degradation of Starch
Starch and starch derivatives have a growing demand in the industry because it is a
renewable and biodegradable resource, abundant, environmentally friendly, cost
competitive, and versatile. Storage starch is deposited in the endosperm (in
amyloplasts) of the seed, while transient starch is deposited in leaves (in chloroplasts).
Starch is deposited as semicrystalline granules and consists of two main glucan
polymers named amylose and amylopectin. Amylose is essentially a linear polymer
with - (1,4) linkages between glucose units, amylopectin is highly branched polyglucan.
In amylopectin molecules, linear chains connecting glucose monomers via -(1,4)-
linkages, are interconnected via -(1,6) linkages. The starch granule is mainly (98- 99%)
composed of amylose and amylopectin, as discussed above. Although, the ratio of
amylose to amylopectin in a normal starch is about 1:3, the exact value depends on the
botanical origin.
Synthesis of Starch involves the simultaneous synthesis of amylase (with - (1,4)
linkages) and amylopectin (with -(1,6) linkages)
The biosynthetic steps required for starch biosynthesis are relatively simple, involving
three committed enzymes:
ADP- glucose pyrophosphorylase (ADPGPase)
starch synthase (SS)
starch branching enzyme (SBE)
Both amylose and amylopectin are synthesized from ADP- glucose, which is
synthesized from glucose-1-phosphate and ATP in a reaction that is catalyzed by
ADPGPPase and that liberates pyrophosphate.
This enzyme is active within the plastid, which means that its substrates, glucose-1-
phosphate and ATP, must also be present in the plastid. In chloroplasts, ATP may be
derived from photosynthesis, but in nonphotosynthetic plastids, it must be specifically
imported from the cytosol, probably by an ADP/ATP translocator.
The glucose-1-phosphate can be supplied by the reductive pentose phosphate
pathway in chloroplasts via phosphoglucoisomerase and phosphoglucomutase.
2. In the next step of starch synthesis, SS catalyzes the synthesis of an a(1-4) linkage
between the nonreducing end of a preexisting glucan chain and the glucosyl moiety of
ADP- glucose, causing the release of ADP. SSs can use both amylose and amylopectin
as substrates in vitro
3. The (1-6) branches in starch polymers are made by SBE, which hydrolyzes an a(1-
4) linkage within a chain and then catalyzes the formation of an a(1-6) linkage between
the reducing end of the "cut" glucan chain and another glucose residue, probably one
from the hydrolyzed chain. Branches are not created randomly, as discussed
previously, but show an aver- age periodicity of 20 glucan residues. SBEs show some
specificity for the length of the a(1-4)glucan chain that they will use as a substrate. Part
of this selectivity may reside in the fact that these enzymes cleave only those glucan
chains that are in a stable double helical conformation, a structure that requires a
minimum glucan chain length


Breakdown of Starch constituents
Breakdown of Starch to yield its constituent D Glucose units may take place in two ways:
1. By the enzyme Diastase
Diastase
Starch Glucose
+H
2
O
In fact diastase is not a single enzyme but a complex of many enzymes which are as
following:
i) - amylase
ii) - amylase
iii) R- Enzyme
iv) Maltase
- amylase attack 1:4 linkage while R-enzyme attack 1:6 Linkage so that the starch is
hydrolyzed to yield Disaccharide units maltose, finally maltase converts maltose into
glucose.
2. By the enzyme Starch phosphorylase
Starch
Phosphorylase

Starch+ Phosphate Glucose-1- Phosphate

Phosphatse converts Glucose 1 phosphate in Glucose.

Synthesis and degradation of Fructans

Fructans are probably the most abundant storage carbohydrate in plants next to starch
and sucrose. Fructans are linear or branched polymers of mostly s-fructosyl-fructose
linkages. Unlike sucrose they are synthesized and stored in vacuoles and can
accumulate in the stems, bulbs and tubers of a number of plants. The basic structure of
a fructan is a trisaccharide, known as a kestose , which has a sucrose molecule linked
to one additional fructose. They are found as oligosaccharides of 5-10 fructose
molecules, ~50 residues in the inulin and ~200 for the levan. Branched fructans found
in grasses, known as graminanes.
Fructan synthesis occurs in vacuoles and sucrose is the precursor.In the first step of
fructan synthesis, the fructose moiety of a sucrose molecule is transferred to a second
sucrose molecule by sucrose-sucrose fructosyl transferase forming a kestose +glucose
Fructose groups are preferentially transferred from a trisaccharide kestose to
longer kestoses rather than from sucrose by fructan-fructan fructosyl
transferases



Degradation of Fructans

Fructan exohydrolases degrade fructan polymers by cleaving terminal fructose
residues.










Synthesis of Cellulose
Cellulose is one of the most important primary plant products. A distinguishing feature of
plant cells is the presence of a cell wall, which is required not only for its structural traits but also
for its many functional properties. Plant cell walls are not homogeneous - they are a complex
mixture of polysaccharides and proteins.
Cellulose consists of unbranched polymers of -linked glucose residues arranged in linear
chains, where every other glucose residue is rotated approximately 180. And hence the
cellulose is an aggregate of -1,4- linked glucan chains. The length, number and organization of
glucan chains in cellulose can be highly variable depending on the source from which it is
obtained.
In broad terms, the synthesis of cellulose can be divided into a polymerization step (where the
glucose residues are joined to form glucan chains) and a post- polymerization step (where the
individual glucan chains associate to form crystalline cellulose and other higher order structures).
Cellobiose is the structural repeating unit of the glucan chains in cellulose. This is in contrast to
other glucan polymers such as starch (alpha-1,4-glucan) and callose ( -1,3-glucan) where
glucose is the repeating unit.