“…Antioxidant system includes antioxidant enzymes (superoxide dismutase: SOD, ascorbate peroxidase: APX, glutathione reductase: GR, catalase: CAT, and guaiacol peroxidase: GPX) and non-enzymatic antioxidants (ascorbic acid: AsA and glutathione: GSH), which can alleviate oxidative damage of biomacromolecules (proteins, lipids, DNA, and RNA) and biomembrane by scavenging excess ROS and MG; osmotic adjustment substances is also known as osmolytes or compatible solutes ( Ashraf and Foolad, 2007 ; Jha et al, 2014 ; Chaudhuri et al, 2017 ; John et al, 2017 ; Li et al, 2017a , b ). Osmotic adjustment substances have emerged multidimensional functions, implicating in osmotic adjustment (to combat with osmotic stress), redox buffering, and ROS scavenging (to ameliorate oxidative stress), and acting as small molecular chaperones (to stabilize the structures of protein, enzyme, DNA, RNA, and biomembrane) and development signaling molecule ( Chen and Murata, 2011 ; Ahmad et al, 2013 ; Chaudhuri et al, 2017 ; John et al, 2017 ). In higher plants, Δ 1 -pyrroline-5-carboxylate synthetase (P5CS), ornithine aminotransferase (OAT), betaine aldehyde dehydrogenase (BADH), and trehalose-6-phosphate phosphatase (TPP) are the rate-limiting enzymes in Pro, GB, and Tre biosynthesis, respectively.…”