Exogenous 5-Aminolevulinic Acid Promotes Osmotic Stress Tolerance of Walnuts by Modulating Photosynthesis, Osmotic Adjustment and Antioxidant Systems

The walnut (Juglans regia L.) is an important economic tree worldwide, often cultivated in arid and semiarid regions. Improving the drought tolerance is of significance for walnut growth, production, and economic effectiveness. 5-Aminolevulinic acid (ALA) is a novel plant growth regulator which raises plant tolerance to various stressful conditions. Here, foliage application of ALA was carried out to uncover its effect on walnuts under polyethylene glycol (PEG) 6000-stimulated osmotic stress. Our results displayed that exogenous ALA greatly promoted the chlorophyll content, photochemical activities, and gas exchange in walnuts under osmotic stress. ALA led to a considerable accumulation of compatible osmotic solutes, enabling walnuts to maintain osmotic equilibrium against drought stress. Furthermore, ALA alleviated the reactive oxygen species (ROS) damages on osmotically stressed walnuts through enhancing the antioxidant enzyme activities, as well as decreasing the ROS and malondialdehyde (MDA) content. The relative water content (RWC) in the ALA-treated leaves was higher than that of PEG-stressed, while the RWC in the substrate of ALA treatment was significantly lower than that of the PEG-stressed, further suggesting that ALA promotes plant water uptake from the substrate under osmotic stress. These demonstrate that ALA improves the photosynthesis, osmotic adjustment, antioxidant systems and, consequently, the walnuts' drought tolerance.