Putrescine enhances chilling tolerance of tomato (Lycopersicon esculentum Mill.) through modulating antioxidant systems

Seedlings of two tomato (Lycopersicon esculentum Mill.) cultivars, cv. Mawa (chilling-resistant) and cv. Moneymaker (chilling-sensitive) were used to investigate the effects of exogenous putrescine (Put) on chilling tolerance as well as on changes of physiological features and the fluctuation of free and conjugated endogenous polyamines (PAs) contents in the leaves under chilling stress. During chilling stress, accumulation of hydrogen peroxide (H2O2) was obviously detected in the leaves of both cultivars, but it was fewer in cv. Mawa. Meanwhile, d-arginine (d-Arg), a Put biosynthesis inhibitor caused more H2O2 accumulation in both cultivars, especially in cv. Moneymaker. By adding back Put to leaves, accumulation of H2O2 obviously reduced in two cultivars. Put was also involved in the increase of Fv/Fm and the decrease of malondialdehyde (MDA) in two cultivars under chilling stress. Despite the two cultivars displaying differential behavior towards enzymic antioxidants, enzymes and components of the ascorbate-glutathione (AsA-GSH) cycle in responses to chilling stress, d-Arg treatment diminished the enzyme activities and antioxidant contents induced by chilling stress and its reversion was performed by adding Put in both cultivars. During chilling stress, free and conjugated endogenous PA contents increased in two cultivars. d-Arg treatment inhibited the increases, and exogenously applied Put enhanced the increases in two cultivars. These results suggested that Put played important roles in the tolerance of tomato against chilling stress, which was most likely achieved by modulating antioxidant system as well as increasing free and conjugated PAs.