Water stress memory in wheat/maize intercropping regulated photosynthetic and antioxidative responses under rainfed conditions

Drought is a most prevalent environmental stress affecting the productivity of rainfed wheat and maize in the semiarid Loess Plateau of China. Sustainable agricultural practices such as intercropping are important for enhancing crop performance in terms of better physiological and biochemical characteristics under drought conditions. Enzymatic and non-enzymatic antioxidant enzyme activities are associated with improved abiotic tolerance in crop plants, however, its molecular mechanism remains obscure. A 2-year field study was conducted to evaluate the influence of intercropping treatment viz. wheat mono-crop (WMC), maize mono-crop (MMC), intercropping maize (IM) and wheat (IW) crops, and nitrogen (N) application rates viz. control and full-dose of N (basal application at 150 and 235 kg ha^-1 for wheat and maize, respectively) on chlorophyll fluorescence, gas exchange traits, lipid peroxidation, antioxidative properties and expression patterns of six tolerance genes in both crops under rainfed conditions. As compared with their respective monocropping treatments, IW and IM increased the Fo/Fm by 18.35 and 14.33%, PS-11 efficiency by 7.90 and 13.44%, photosynthesis by 14.31 and 23.97%, C-capacity by 32.05 and 12.92%, and stomatal conductance by 41.40 and 89.95% under without- and with-N application, respectively. The reductions in instantaneous- and intrinsic-water use efficiency and MDA content in the range of 8.76-26.30% were recorded for IW and IM treatments compared with WMC and MMC, respectively. Compared with the WMC and MMC, IW and IM also triggered better antioxidant activities under both N rates. Moreover, we also noted that intercropping and N addition regulated the transcript levels of six genes encoding non-enzymatic antioxidants cycle enzymes. The better performance of intercropping treatments i.e., IW and IM were also associated with improved osmolytes accumulation under rainfed conditions. As compared with control, N addition significantly improved the chlorophyll fluorescence, gas exchange traits, lipid peroxidation, and antioxidant enzyme activities under all intercropping treatments. Our results increase our understanding of the physiological, biochemical, and molecular mechanisms of intercropping-induced water stress tolerance in wheat and maize crops.