Week 9

CARBOHYDRATE
METABOLISM IN
PLANT
The link source to sink

Materials are translocated in the phloem
from sources (areas of photosynthate
supply) to sinks (areas of metabolism or
storage of photosynthate).

Sources are usually mature leaves.

Sinks include organs such as roots and
immature leaves and fruits.

Carbohydrates are stored in the leaves as starch and exported as
sucrose to other parts of the plants such as the roots or growing
seeds

CO2 assimilation in the chloroplasts yields triose phosphate, which is
transported by the triose phosphate-phosphate translocator
The compartments of carbohydrate
metabolism
Synthesis of starch and sucrose
Synthesis of starch Synthesis of sucrose

Starch is synthesis in
In cytosol
the chloroplas
Substrate: triose

Substrate: triose phosphate
phosphate
When the cytosolic

When the cytosolic Pi concentration is
Pi concentration is high
low
Sucrose is synthezid

Starch is synthesized via UDP-glucose
via ADP-glucose
Synthesis of fructans

Fructans are
In fructans of the 6-
deposited as storage kestose type, the
substances in the chain consists of n =
vacuole 10 to 200 and in

Fructans are derived those of the 1-
from kestoses. kestose type of n <

They are formed by
50 fructose residues.
the linkage of
In fructans of the
fructose residues to neokestose type, n
a sucrose molecule. and m are < 10.

Sucrose is precursor for synthesis of kestoses.

Three important reactions of the kestose
biosynthesis pathway proceeding in the vacuole
are shown

The conversion of CO2
assimilate to fructan.

Fructose 6-phosphate,
which is provided as a
product of
photosynthesis in the
cytosol, is first converted
to sucrose.

The glucose required for this is formed as a by-product in

the synthesis of fructan in the vacuole.

Phosphorylation is catalyzed by a hexokinase present in
the cytosol.

The entry of sucrose into the vacuole and the release of
glucose are facilitated by different translocators.
The carbohydrate of cell wall

Cellulose and callose
Synthesis of cellulose

Location in the plasma membrane

Catalyzed by cellulose synthase

Substrate UDP-glucose from cytosol
The glyoxylate cycle

The glyoxylate cycle is active in the germinating seeds of
some plants and in certain microorganisms that can live
on acetate as the sole carbon source.

In plants, the pathway takes place in glyoxysomes in
seedlings.

It involves several citric acid cycle enzymes and two
additional enzymes: isocitrate lyase and malate synthase.
Reaction of glyoxylate cycle
Relationship between the
glyoxylate and citric acid
cycles
Coordinated regulation of glyoxylate
and citric acid cycle
Week 9

LIPIDS METABOLISM IN
PLANT
Synthesis of fatty acids

Location: plastid
in the chloroplasts of green cells
in the leucoplasts and chromoplasts of non-green
cells.

also found in the membrane of the ER, these
enzymes appear to be involved only in the
modification of fatty acids
chain elongation of fatty acids, as catalyzed by
elongases,
the introduction of further double bonds by
desaturase

Acetyl CoA is the precursor for the synthesis
of fatty acids

Acetyl CoA can be formed from Acetate and
Pyruvat

Reaction sequence
for the synthesis of
fatty acids:
activation,
condensation,
Reduction, release of
water,
And further reduction
elongate a fatty acid by
two carbon atoms.

Acetyl CoA carboxylase reaction scheme.

The biotin linked to the biotin carboxyl carrier protein reacts in
turn with biotin carboxylase and carboxyl transferase.

The circular motion has been chosen for the sake of clarity; in
reality, there probably is a pendulum-like movement.

The eukaryotic form of acetyl CoA carboxylase is present as a
multifunctional protein.

Biotin is linked via a lysine residue to the biotin
carboxyl carrier protein.
 Further steps of fatty acid synthesis are also catalyzed
by a multienzyme complex

The acyl carrier protein (ACP), located in the center, carries the
fatty acid residue, bound as thioester, from enzyme to enzyme.

The circular movement has been chosen for the sake of clarity, but
it does not represent reality.

The acyl carrier protein (ACP) contains pantetheine,
the same functional group as coenzyme-A.

Elongation and
desaturation of fatty
acids in plastids.

Stearoyl-ACP desaturase, localized in the
plastids,
catalyzes the desaturation of stearoyl-ACP to oleyl-
ACP.
The reaction can be regarded as a monooxygenation
with the subsequent release of water.

Acyl lipid desaturases, integral proteins of the ER
membrane,
Catalyze the desaturation of fatty acids, which are parts
of phospholipids.
Electron transport from NADPH to the desaturase
requires two more proteins, one NADPH-Cyt-b and
cytochrom-b5.

Acyl-ACPthioesterases release mainly 16:0 and 18:1-
fatty acids and only low amounts of 18:0-fatty acids.

After the fatty acids leave the stroma and enter the
cytosol, they are immediately converted to acyl-CoA.

Acyl-ACP formed in the plastids has two
important functions:
1. It acts as an acyl-donor for the synthesis of
plastid membrane lipids.
2. For biosynthesis outside the plastids, acyl-
ACP is hydrolyzed by acyl- ACP
thioesterases to free fatty acids, which then
leave the plastids
The synthesis of lipid membrane
Overview of the synthesis of membrane lipids.
Formation of CDP-choline
The membrane lipids synthesized in the plastids and at
the ER have different fatty acid compositions.
The ER membrane is the site of fatty acid elongation
and desaturation
Some of the plastid membrane lipids are formed
via the eukaryotic pathway

Part of the membrane lipids of the

thylakoid membranes are
synthesized via the eukaryotic
pathway (red) and other parts
via the prokaryotic pathway
(black).
The Synthesis of Triacylglycerols
Triacylglycerols are formed in the membranes of
the endoplasmic reticulum
Triacylglycerols are storage substances

An oil body enclosed by a lipid monolayer is formed by the

incorporation of triacylglycerols in an ER membrane.
Oleosins are anchored to the lipid monolayer
Lipid Mobilization

During seed germination, storage lipids
are mobilized for the production of
carbohydrates in the glyoxysomes
Utilization of lipid from plant
1. Plant fat is used for nutrition