BIOSYNTHESIS OF LIPIDS 607

BIOSYNTHESIS OF LONG-CHAIN FATTY

ACIDS
(=M TOCHONDRIAL AND MICROSOMAL FATTY ACID SYNTHESIS)
Palm.
tale (a C16 fatty acid) i1s the
major product of the fatty acid
also as the de novo
calledas
system in that the synthase system. This system
lso
andmalony/-Co4 palmitate is
CoA (ACP). In plants and animals, the most constructed from acetyl-CoA (ACP
and linolenic. Theseimportant fatty
namely stearic, oleic, linoleic acids are the Cia tatty
CIds,

stems which differ Ci8 fatty acids are synthesized by

e l o n g a i o ns Y s l e m
markedly from the de novo system. Palmitate acts as a
ano-chain fatty acids (Fig. 25-12). It may be lengthened to form
precursor
nated fatty acids by further addition of stearate (18:0) or even
acetyl groups
l a g e rs a t u r a

present in the
the action of through fatty acid
elongatie system
endoplasmic reticulum (= microsomes) and mitochondria in the
animals and in the soluble cytosol in the case of
of plants.
case
Palmitate
(16:0)
Desaturation
Elongation
Palmitoleate
16:1(49)
Stearate Elongation
18:0
Vaccenate
Elongation 18:1(A)
Desaturation
Longer, saturated
Oleate fatty acids
18:1(4)
Desaturation
(in plants only)
Linoleate
18:2(49.12)
Desaturation
Desaturation
(in plants only)

a-linolenate y-linolenate
18:3(A9.12,15 18:3(A6.9,12)
Elongation

Other polyunsaturated Eicosatrienoate

fatty acids 20:3(A8.11,14)

Desaturation

Eicosatetraenoate
(=Arachidonate)
20:4(458.1,14)
Fig. 25-12. Biosynthesis of some fatty acids in mammals
ralhymitate is the
precursor
precursor of
acids, palmitoleate ol stearate and longer-chain saturated fatty acids, as well as the
monounsaturated

sqired i
ads the diet as
and oleate. Mammals cannot convert oleate into linoleate or a-linolenate, hence
and essential acids (EFAs). Conversion of linoleate into other polyunsaturated fatty
eicoandnoids is also fatty
putlined.
 608 FUNDAMENTALS OF BIOCHEMISTRY
In animals
Endoplasmic reticulum membrane
Malonyl-CoA Stearyl-CoA
Palmitoyl-CoA + NADPH

Mitochondrial outer and inner membranes

Acetyl-Co0A
Stearyl-CoA
Palmitoyl-CoA + NADPH

In plants:
Soluble cytosolic system
Malonyl-ACP
Stearyl-ACP
Palmitoyl-ACP
+NADPH

involved and coenzyme A, rather than ACP,

the acv
is
are
Although different enzyme systems the mechanism of elongation is otherwie
carrier directly involved in animal fatty
acid synthesis,
donation of two carbons by malonyl-AC?,
identical with thatemployed in palmitate synthesis: reduction (with NADPH) tothe
and another
followed by reduction (with NADH), dehydration
largely abolishes chain elongation. Elongationo
saturated CIR product, stearyl-CoA. Fasting
myelination in order to provide C and C faty
stearyl-CoA in brain increases rapidly during
acids that are present in sphingolipids.