Transaminationreactions involve the transfer of an amino group from one
compound to another. For instance, asparagine synthase(AS;Fig. 2) converts
aspartate and glutamine to asparagine and glutamate. Asparagine contains a
high quantity of nitrogen (2N) for each carbon (4C) present, compared with
glutamate (1N:5C) or glutamine (2N:5C). Expression of the AS gene is repressed
(inhibited) by high light and high carbohydrate production, so the plant makes
nitrogen compounds with low quantities of nitrogen. When light is low and
carbohydrate is scarce, carbon is conserved by the synthesis of compounds with
a much higher nitrogen content.
Plants synthesize a wide variety of lipids, including: membrane lipids, in which
long-chain fatty acids, commonly 16–18 carbons in length, are esterified to
glycerol; the waxes of the cuticle and suberin of the endodermis; and the
storage lipids of seeds (most commonly triacylglycerols).
Plants use acetyl CoA as the basic building block for assembling long-chain
fatty acids. Acetyl CoA is produced by pyruvate dehydrogenase present both
in plastids and mitochondria. In the chloroplast, pyruvateis produced from
3-phosphoglycerate from the Calvin cycle; in non-photosynthetic tissue, it origi-
nates from glycolysis. Synthesis of glycerolipids occurs in two stages: addition
of the fatty acid chains to glycerol-3-phosphate and the addition of a head
group. Glycerol-3-phosphate is derived in stages originating from glyceralde-
hyde-3-phosphate. In photosynthetic tissue, the chloroplast is the chief source of
fatty acids, which may be used directly in the chloroplast or exported to the
cytoplasm. Plant cells can synthesize glycerolipids at the endoplasmic
reticulum (ER) and in mitochondria as well as in plastids; all three of these
organelles are believed to be inter-connected and lipids traffic between them
and to the other cellular membranes.
Seeds make triacylglycerols, which are storage oils, and frequently very
important human foods. Up to 60% of the dry weight of a seed may be in the
form of storage oils. Triacylglycerols consist of a glycerol to which three fatty
acid chains have been added (Fig. 3). They are stored in oil bodies, small lipid
droplets with a surrounding coating of lipid and protein. The oil is first synthe-
sized in the ER.
Lipid
biosynthesis
156 Section J – Metabolism
Glutamate Oxaloacetate Aspartate 2-oxoglutarate
Glutamine ++Aspartate Asparagine Glutamate
COOH
HC NH 2
CH 2
CH 2
CO–
O
NH 2
COOH
CH 2
CO–
O
C O
COOH
CH 2
CO–
O
CH 2
C O
COOH
CH 2
CO–
O
C
++
AAT
Aspartate
aminotransferase
AS
Asparagine
Synthetase
Fig. 2. Other amino acids are synthesized by transamination reactions which occur in various
organelles.