Abstract
This review focuses on the biochemical regulation and function of the alternative
oxidase in vivo. About 10 years ago, two activation mechanisms were discovered in isolated mitochondria,
namely activation by reducing sulfur bonds in the protein and activation by an allosteric
effect of pyruvate. It was proposed that plants would have a regulatory mechanism
to modify alternative oxidase activity in vivo. However, more recent studies have shown that these two activation mechanisms may
not play such an important role in regulation of alternative oxidase activity in vivo after all. Pyruvate and reduction of the sulfide bonds in the protein are definitely
required for alternative oxidase activity, but they do not appear to be regulating
the activity in vivo.
Despite the energy wasting nature of the alternative oxidase, there was no obvious
physiological function for the pathway for many years. It is now more clear that the
alternative oxidase can prevent the production of excess reactive oxygen species radicals
by stabilizing the redox state of the mitochondrial ubiquinone pool, while allowing
continued activity of the citric acid cycle. This may be important under conditions
when the NADH supply is relatively high (reductant overflow), or when the cytochrome
pathway is restrained. The cytochrome pathway might be inhibited by naturally occurring
cyanide, nitric oxide, sulfide, high concentrations of CO2 , low temperatures, or by limited phosphate supply.
Abbreviations
AOX: alternative oxidase
BHAM: benzhydroxamic acid
SHAM: salicylhyroxamic acid
Q: ubiquinone
Qox : oxidized ubiquinone
Qr : reduced ubiquinone
Qt : total pool of ubiquinone
Qr /Qt : relative reduction state of the ubiquinone pool
Key words
Alternative oxidase - respiration - ubiquinone - oxygen free radicals - pyruvate
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F. F. Millenaar
Plant Ecophysiology Utrecht University
Sorbonnelaan 16 3584 CA Utrecht The Netherlands
eMail: f.f.millenaar@bio.uu.nl
Section Editor: L. A. C. J. Voesenek