It is well known that antioxidants such as AA (reduced ascorbate), glutathione (GSH) (reduced glutathione) and melatonin can delay seed ageing. Can they recover aged seed? Artificial aged maize seeds were obtained and their reduced germination rate (GR) and high lipid peroxidation were recorded. Exogenous melatonin was applied on these aged seeds and enhanced GR was observed. However, treatment with other antioxidants such as AA, GSH or DMTU (dimethyl thiourea) did not significantly improve or even reduce the GR of aged seeds. In addition, melatonin improved germination ability of theses aged seeds can be significantly impaired by DDC (diethyldithiocarbamic acid, a specific inhibitor of superoxide dismutase or superoxide dismutase (SOD)) and ATZ (aminotriazol, a specific inhibitor of catalase or CAT). In a further study, we found that melatonin but not other antioxidants (AA, GSH and DMTU) significantly induced CAT and SOD activities of aged seeds after imbibition. Accordingly, melatonin significantly reduced lipid peroxidation in aged seeds than that of other antioxidants. Taken together, these data suggest that melatonin induced antioxidant enzyme but not its direct reactive oxygen species (ROS) scavenging capacity contributing to recovery of aged maize seeds.

The role of exogenous adenosine 5'-triphosphate (ATP) in the regulation of antioxidant response in plants under heavy metal stress is unclear. Here, we investigated the effects of exogenous ATP application on plant growth, antioxidant response, and Cd accumulation in maize seedlings. Treatment with 0.1 mM CdCl₂ moderately reduced dry weight, decreased chlorophyll content, impaired photosynthesis, and increased lipid peroxidation in maize seedlings compared with controls. However, toxicity due to Cd was alleviated after 10-200 µM ATP treatment. Subsequently, the activity of Cd-regulated antioxidant enzymes, antioxidant metabolite accumulation, and total antioxidant capacity were drastically enhanced after 50 µM ATP treatment. Similar patterns were observed in the ADP-treated group but not in the AMP-treated group under Cd stress. However, the ATP-induced elevation in antioxidant defense ability was decreased by the inhibition of NADPH oxidase (NOX). ATP-induced elevation in NOX activity and H₂O₂ production was partly reversed by the inhibition of NOX in maize seedlings under Cd stress. Furthermore, ATP promoted Cd accumulation in the roots and shoots of maize seedlings. However, the ATP-induced increase in Cd accumulation was partly abolished by the inhibition of NOX. To our knowledge, this is the first report on the role and mechanism of exogenous ATP in regulating plant growth, antioxidant response, and heavy metal phytoextraction. The study provides a new method based on exogenous ATP for enhancing heavy metal tolerance in plants.