Biosynthesis of Storage Carbohydrates

(Starch)

Jeeva.V
2023533004
M.Sc,. Horticulture
 Introduction
• Photosynthesizing cells of plants make and store starch in
their chloroplasts to meet their metabolic needs.
• translocate to regions of plants that are not
photosynthetically competent
• Sinks – roots, seeds and developing leaves
• Sucrose – the form in which products of carbohydrate
catabolism are transported to developing seedlings.
• Excess glucose is converted into polymeric forms for storage
and transport in wide range of organisms.
• Principal storage forms of glucose in plants
• Starch
• Inulin
• Transport form
• Sucrose in plants or its galactosylated derivatives.
• The starch can be converted to sucrose and exported at night.
Interconversion of sucrose and starch
 important aspect in carbohydrate metabolism in plants
 need to maintain the continuous supply of sucrose at all times during
the diurnal cycle in photosynthetic tissues
 exported from the leaf to provide the carbon source for respiration
and growth of the whole plant.
• Sucrose synthesis and starch synthesis is regulated based on the 3PGA/Pi
level in the chloroplasts.
• Precursor for sucrose synthesis is Triose PO4 derived from calvin cycle
during the day or derived from starch breakdown at night
• Triose PO4 required for sucrose formation are exported from stroma of
chloroplast.
• Sucrose synthesis occurs in the cytoplasm and is transported to other
parts.
• Excess carbon during photosynthesis is stored as starch in the chloroplasts.
Triose phosphates produced in the Calvin cycle can
be used for starch or sucrose synthesis

chloroplast
Calvin
cycle
Starch

Triose- P

Sucrose
cytosol
 Starch

• Principal food-reserve polysaccharide composed of alpha-glucan

• occurs in the seeds, fruits, leaves, tubers and bulbs - few percent to over
74%.
• Components: Amylose and Amylopectin
• 2% of amylose in waxy rice or waxy maize to about 80% amylose in
amylomaize.
• majority of starches - 15 to 35% of amylose.
• ratio of amylose and amylopectin is a function of the enzymes
• Granule bound starch synthase (GBSS) helps in the synthesis of
Amylose.
• Soluble starch synthase (SSS) helps in the synthesis of Amylopectin.
• Starch granules consist primarily of amylopectin, and amylose.
• Starch granules contain the enzymes for starch synthesis and degradation.
• These enzymes are present in several isoforms, some of which are bound
to the starch granules whereas others are soluble.
• Amylose consists mainly of unbranched chains of about 1,000 glucose
molecules.
• Amylopectin, with 104 to 105 glucose molecules, is much larger than
amylose and has a branching point at every 20 to 25 glucose residues.
• Results of X-ray structure analysis show that a starch granule is
constructed of concentric layers.
 CH 2 OH 6 CH OH CH 2 OH CH 2 OH CH 2 OH
2
O 5 O H O O H H O H
H H H H H H
H H H H H
OH H 1 4 OH H 1 OH H OH H OH H
O O O O OH
OH 2
3
H OH H OH H OH H OH H OH
am ylose

CH 2OH CH 2OH
H O H H O H amylopectin
H H
OH H OH H 1
O
OH
O
H OH H OH

CH 2OH CH 2OH 6 CH 2 CH 2OH CH 2OH

H O H H O H H 5 O H H O H H O H
H H H H H
OH H OH H 4 OH H 1 4 OH H OH H
O O O O OH
OH
3 2
H OH H OH H OH H OH H OH

Starch – amylose & amylopectin

 Starch Granules
Starch granules are classified as transitory or reserve.

Transitory starch granules accumulate for only a short period of time before
they are degraded, e.g.

- Starch formed in leaf chloroplasts during the day and hydrolyzed and
transported to other plant parts at night as simple sugar.

Reserve starch, an energy storage for germination is formed in amyloplasts.

- In cereals the reserve starch often represents 65% to 75% and in potato
 Starch metabolism
• Starch synthesis
• enzymes for starch synthesis are localized in Plastids chloroplast and amyloplast.
• Starch breakdown
• two possible pathways for mobilization or catabolism of starch.
• Hydrolytic process
• Amylases convert starch to maltose, maltotriose and dextrins
• further converted to glucose by ‑ glucosidase.
• Glucose is then phosphorylated to glucose 6-phosphate by hexokinase.
• This is glycolytically converted to triose phosphate
• Phosphorolytic process
• starch can be directly degraded to glucose 1-phosphate by starch
phosphorylase
• Glucose 1-phosphate is then converted to glucose 6-phosphate by
phosphoglucomutase.
 Starch Synthesis in Chloroplast and
Amyloplast
•Enzymes for starch synthesis are localized in chloroplast and
amyloplast.
•Starch synthesis and storage takes place in chloroplasts, a type of
pigmented plastid involved in photosynthesis.
•Amyloplasts are non-pigmented organelles found in some plant
cells. They are also responsible for the synthesis and storage of
starch granules, through the polymerization of glucose.
Starch is made in photosynthetic and
non-photosynthetic cells

Photosynthetic cells
 transitory starch
storage
 green leaves

Non-photosynthetic cells: Sucrose

 long-term reserve starch
storage. Starch
 roots, tubers, seeds .
Starch synthesis
Photosynthesis Fructose 6 P

Glucose – 1 P + ATP
ADP - Glucose pyro
phosphorylase

ADP Glucose + PPi

Primer Starch synthase

Starch
 Biosynthesis of Starch
• Biosynthesis of starch takes place in Chloroplast and Amyloplast.
• Glucose -6 - Phosphate converted to Glucose -1-phosphate with the help of an enzyme
Phospho glucomutase.
• ATP is added to Glucose -1- Phosphate in the presence of an enzyme ATP-Glucose
Pyrophosphorylase to form ADP-Glucose.
• A primer is added to ADP-Glucose (Glucosyl donor) in the presence of an enzyme Starch
synthase to form amylose.
• In presence of branching enzyme amylopectin is formed.
• Branches lengthen by Starch synthase.
ATP + Glc -1-Phosphate ADP-Glc + PPi
nADP-Glc + Primer Amylose + n ADP
Amylose + nADP-Glc Amylopectin
• Soluble Starch Synthase (SSS) helps in the synthesis of Amylopectin.
• Granule Bond Starch Synthase (GBSS) helps in the synthesis of
Amylose.
• Starch in amyloplast is mobilised & broken down to Glucose and
Glucose-1-Phosphate by amylases and starch phosphorylase.
• Glucose and Glucose-1-Phosphate is converted to Glucose-6-
Phosphate and transported to cytoplasm and converted to Sucrose.
• Branches are formed by a branching enzyme.
• At certain chain lengths, the polysaccharide chain is cleaved at the
glycosidic bond (alpha 1→4) and the polysaccharide fragment thus
separated is connected via a newly formed (alpha 1→6) bond to a
neighboring chain.
• These chains are elongated further by starch synthase until a new
branch develops.
• In the course of starch synthesis, branches are also cleaved again by a
debranching enzyme.
• It is assumed that the activities of the branching and the debranching
enzymes determine the degree of branching in starch.
ADP-Glucose Is the Substrate for Starch Synthesis in Plants
ADP-glucose pyrophosphorylase
• AGPase brings about the first committed step in the biosynthetic
pathway leading to starch production in all the plants.
• AGPase is a heterotetramer of 2 large (54-60Kd) and 2 small (51-
55 Kd) subunits.
• Both subunits are required for the activity.
• Small subunit is thought to be main catalytic activity, large
subunit is regulatory.
• Generally, this enzyme is allosterically regulated by 3-
phosphoglycerate (activator) and inorganic orthophosphate
(inhibitor).
Starch biosynthesis is regulated by
ADP-glucose pyrophosphorylase
Starch biosynthesis by starch synthase
Starch synthase
 Starch Synthase(SS) catalyzes a 1,4- linkage between
nonreducing end of glucan chain & Glc from ADP-Glc.

 SS can use both amylose and amylopectin as acceptors.

Starch branching enzyme (SBE)

 Starch branching enzyme (SBE) catalyzes a 1,6- linkage
forming side chains.