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Journal of Lipid Research, Vol 36, 2471-2477, Copyright © 1995 by Lipid Research, Inc.
H Sprecher, DL Luthria, BS Mohammed and SP Baykousheva
Recent studies refute the commonly accepted, but untested, hypothesis that
7,10,13,16-22:4 and 7,10,13,16,19-22:5 are desaturated at position 4 by a
microsomal acyl-CoA-dependent desaturase. The synthesis of
4,7,10,13,16,19-22:6 occurs via the following reaction sequence:
7,10,13,16,19-22:5-->9,12,15,18,21-24:5-->6,9,12,15,18,21-24:6
4,7,10,13,16,19-22:6. The synthesis of 4,7,10,13,16-22:5 from
7,10,13,16-22:4 takes place via an analogous pathway. According to these
pathways the 24-carbon acids that are made in the endoplasmic reticulum
move to a site for partial beta-oxidation, which is most likely
peroxisomes. The products of partial beta-oxidation, 4,7,10,13,16-22:5 and
4,7,10,13,16,19-22:6, then move back to the endoplasmic reticulum where
they are used as substrates for membrane lipid biosynthesis. The ability of
a fatty acid to serve as a substrate for continued peroxisomal
beta-oxidation, versus its transfer out of peroxisomes for subsequent
endoplasmic reticulum-associated esterification reactions, may be an
important control for regulating membrane lipid fatty acid composition.
Indeed, the revised pathways of polyunsaturated fatty acid biosynthesis
imply that there is considerable intracellular movement and recycling of
fatty acids between peroxisomes and the endoplasmic reticulum. In addition,
these revised pathways require that two 18-carbon and two 24-carbon acids
are substrates for desaturation at position 6. Also, as linoleate and
linolenate are metabolized, respectively, to 6,9,12,15,18-24:5 and
6,9,12,15,18,21-24:6, three n-6 acids and three n-3 acids are substrates
for malonyl-CoA dependent chain elongation. It remains to be determined how
many microsomal enzymes are required to carry out these reactions and
whether other ancillary enzymes are expressed in tissues whose membrane
lipids accumulate very long-chain polyunsaturated acids with up to 36
carbon atoms.
REVIEWS
Reevaluation of the pathways for the biosynthesis of polyunsaturated fatty acids
Department of Medical Biochemistry, Ohio State University, Columbus 43210, USA.
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