Impact of ozone on monoterpene emissions and evidence for an isoprene-like antioxidant action of monoterpenes emitted by Quercus ilex leaves

Quercus ilex (L.) leaves emit monoterpenes, particularly alpha-pinene, beta-pinene and sabinene. Apart from the monoterpene pools that are stored in specialized structures and have a clear defensive, or attractive role, the function of monoterpenes in Q. ilex leaves is unknown. We tested whether monoterpenes have an antioxidant role, as has previously been found for isoprene in isoprene-emitting leaves. We exposed Q. ilex leaves to either mild and repeated ozone exposure (Experiment 1) or to a single acute ozone exposure (Experiment II) at temperatures ranging between 20 and 32 degreesC. Both ozone treatments rapidly stimulated monoterpene synthesis, but had no effect on photosynthesis and caused no visible damage to leaves maintained at 25, 30 or 32 degreesC. Ozone inhibited both photosynthesis and monoterpene synthesis in leaves maintained at 20 degreesC. To characterize the relationship between monoterpenes and ozone-induced damage, we fed detached leaves fosmidomycin, a selective inhibitor of isoprene synthesis. Fosmidomycin caused rapid and complete inhibition of monoterpene emissions in leaves maintained at 30 degreesC, confirming that monoterpenes are synthesized by the same biochemical pathway as isoprene. However, over the experimental period, fosmidomycin did not affect concentrations of compounds that are formed from chloroplastic isoprenoids and that might have conferred antioxidant protection, either directly (carotenoids) or indirectly (chlorophylls, xanthophylls). In leaves whose monoterpene synthesis had been inhibited by fosmidomycin, ozone rapidly and significantly inhibited photosynthesis and increased the production of hydrogen peroxide and malonyldialdehyde. We conclude that monoterpenes produced by Q. ilex leaves share the same biosynthetic pathway and function as isoprene. Furthermore, all volatile isoprenoids may have similar antioxidant properties and may be stimulated by the same stress-inducing conditions.