Melatonin alleviates iron stress by improving iron homeostasis, antioxidant defense and secondary metabolism in cucumber

Iron is an essential element for plants and animals; however, deficit or excess iron can result in stress on plants. In this study, we unveiled the mechanisms of shoot-based tolerance to iron toxicity by melatonin (N-acetyl-5-methoxytryptamine) in cucumber plants. Both low and high iron (Fe) supply in hydroponics decreased growth and biomass accumulation, induced chlorosis and oxidative stress, and reduced chlorophyll content, photosynthesis rate and transpiration rate in cucumber leaves. Notably, the negative effect of low-Fe was more profound than that of high-Fe treatment. However, exogenous melatonin application alleviated those inhibitions in growth, biochemical and physiological parameters, which entailed an elevation of endogenous melatonin content and a reduction of electrolyte leakage, reactive oxygen species accumulation and lipid peroxidation by improving the activity and transcripts of antioxidant enzymes and secondary metabolism-related enzymes, and concentrations of phenols and flavonoids under low and high iron conditions after melatonin treatment. Analysis of iron content in leaves and roots revealed that melatonin significantly increased the iron content under low-Fe conditions, but it decreased the same under high-Fe conditions. Moreover, melatonin increased the transcript levels of FRO2 and IRT1 under low-Fe, but it decreased those transcripts under high-Fe, suggesting that melatonin plays a dual role in iron uptake under low and high iron conditions. This study expands the stress ameliorative role of melatonin in plants and may have potential implications in agronomic management of crops in low and high iron prone soils.