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THE PLANT CELL, Vol 8, Issue 8 1323-1335, Copyright © 1996 by American Society of Plant Biologists
A Nuclear Gene Encoding Mitochondrial Proline Dehydrogenase, an Enzyme Involved in Proline Metabolism, Is Upregulated by Proline but Downregulated by Dehydration in Arabidopsis
T. Kiyosue, Y. Yoshiba, K. Yamaguchi-Shinozaki and K. Shinozaki
Laboratory of Plant Molecular Biology, Institute of Physical and Chemical Research (RIKEN), Tsukuba Life Science Center, 3-1-1 Koyadai, Tsukuba, Ibaraki 305, Japan
Proline is one of the most common compatible osmolytes in water-stressed
plants. The accumulation of proline in dehydrated plants is caused both by
the activation of proline biosynthesis and by the inactivation of proline
degradation; a decrease in the level of accumulated proline in rehydrated
plants is caused both by the inhibition of proline biosynthesis and by the
activation of proline degradation. The proline biosynthetic pathway has
been well characterized, but the degradation of proline is poorly
understood. Sequence analysis of an Arabidopsis cDNA clone, ERD5 (for early
responsive to dehydration stress), isolated from plants dehydrated for 1
hr, revealed that it encodes a protein with identity to products of the
yeast PUT1 (for proline utilization) gene (23.6% over 364 amino acids) and
the Drosophila sluggish-A gene (34.5% over 255 amino acids). Their gene
products are precursors of proline oxidases (dehydrogenase) (EC 1.5.99.8),
which are the first enzymes involved in the conversion of proline to
glutamic acid. Proline oxidase is localized in mitochondria. RNA gel blot
analysis demonstrated that transcripts of the ERD5 gene were undetectable
when plants had been dehydrated for 10 hr, but large amounts of the
transcript accumulated when plants subsequently were rehydrated. Elevated
levels of the transcript were also found in plants that had been incubated
in a medium that contained proline. Immunologically, we showed that the
product of ERD5 is localized in the mitochondrial fraction and accumulates
in response to proline in cultured cells. Fusion genes for ERD5 and PUT1
complemented a put1 mutant of yeast, allowing put1 to grow with proline as
the source of nitrogen. These results suggest that ERD5 encodes a precursor
of proline dehydrogenase (oxidase), which is regulated at the level of mRNA
accumulation in both dehydrated and rehydrated plants.
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