Most crops of economic importance are vulnerable to environmental abiotic stresses, such as drought, extreme temperatures, and salinity (Khan et al., 2017;Piticar, 2019). Temperature is a key factor that affects plant growth and development. Sub-zero temperatures can damage and even kill crops (Piticar, 2019) due to extracellular ice formation that dehydrates cells and subsequently damages membranes (Nagao et al., 2008;Thomashow, 1999), cells disruption due to fusion between cellular and intercellular membranes can occur while the ice is formed (Takahashi et al., 2018;Thomashow, 1999). Low temperature may induce photoinhibition by producing an imbalance