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of supernatant was collected. Soluble sugar concentration was determined by the sulfuric acid anthrone method ( Yemm and Willis, 1954 ). Proline content. 0.5 g of FW leaves was extracted for free proline content measurements according to Bates et al

cryoprotective properties (such as proline and soluble sugars) and root TTC-reducing activity ( Ristic and Ashworth, 1993 ; Uemura et al., 1995 ; Xin and Browse, 2000 ). A number of genetic and molecular studies on plant response to low temperatures have

room temperature. Samples were then boiled in a water bath for 30 min and EL 2 was measured again. The leaf EL was calculated as follows: EL (%) = (EL 1 /EL 2 ) × 100%. Determination of proline and soluble sugar contents. Proline content was determined

germination liquid medium (50 mL) composed of MS salts, 1.156 μM GA 3 , and 3% sucrose. The cultures were incubated at 25 ± 2 °C in darkness for 15 d and later at continuous light for another 15 d. Proline determination. Free proline content was determined

et al., 2009 ). Various metabolites (such as proline) may accumulate during cold acclimation ( Zhang and Ervin, 2008 ; Zhang et al., 2006 , 2008 ). Proline, an amino acid, functions as osmoprotectant and antioxidant to protect cell membrane during

. (2004) found high expression of citric acid, β-alanine, proline (Pro), tyrosine (Tyr), valine (Val), 1,3-diaminopropane, ethanolamine, putrescine, galactose, glucose, and others, especially Pro, under mild drought stress (6 to 7 d without irrigation) in

addition, MDA and proline production have been used as biochemical markers to characterize plant responses to abiotic and biotic stresses ( Dar et al., 2016 ; Irulappan and Senthil-Kumar, 2018 ). Chlorophyll fluorescence parameters also are affected by

measurements. The water content was measured using the formula: water content = (fresh weight − dry weight)/fresh weight. The proline content was measured according to the method of Bates et al. (1973) . The soluble protein content was measured according to

-watered control and drought stress, respectively. eEF-1α was internal loading control. Proline is highly accumulated in the stress-tolerant species under drought stress ( Türkan et al., 2005 ). Both P5CS and P5CR are important enzymes in the proline

most studied osmotic compounds in plants are proline, glycine betaine, and sugars which act as a mediator and stabilizer ( Verbruggen and Hermans, 2008 ). Meanwhile, abiotic stresses such as drought, salinity, chilling, and heavy metals will lead to