Integrative application of licorice root extract or lipoic acid with fulvic acid improves wheat production and defenses under salt stress conditions

Although different plant extracts and plant growth regulators are used as biostimulants to support plants grown under salt stress conditions, little information is available regarding the use of licorice root extract (LRE) or lipoic acid (LA) as biostimulants. Studies on the application of LRE or LA in combination with fulvic acid (FA) as natural biostimulants have not been performed. Therefore, in this study, two pot experiments were conducted to evaluate the potential effects of LRE (5 g L-1) or LA (0.1 mM) supplemented as a foliar spray in combination with FA (0.2 mg kg(-1) soil) on osmoprotectants and antioxidants, growth characteristics, photosynthetic pigments, nutrient uptake, and yield as well as on the anatomical features of the stems and leaves of wheat plants irrigated with three levels of saline water (0.70, 7.8, and 14.6 dSm(-1)). Moderate (7.8 dSm(-1)) and high (14.6 dSm(-1)) levels of salinity caused a significant (p <= 0.05) increase in the activities of SOD, APX CAT, PDX, and GR as well as in electrolyte leakage, malondialdehyde level, and reactive oxygen species (O-2 center dot- and H2O2) levels compared to those in controls (plants irrigated with tap water). However, the leaf relative water content, membrane stability index, NPK uptake, leaf area, plant height, spike length, straw yield, grain yield, and protein content of wheat grains significantly (p <= 0.05) decreased. Addition of LRE or LA and/or HA to wheat plants under saline stress significantly (p <= 0.05) enhanced their morphological and physio-biochemical characteristics in parallel with increases in the activities of enzymatic antioxidants. Salinity stress altered (p <= 0.05) wheat stem and leaf structures; however, treatment with LRE + FA significantly improved these negative effects. These findings indicate that FA + LRE treatment significantly improved the antioxidant defense system of the plants, thereby reducing ROS levels and increasing wheat growth and production under saline conditions.