Glutathione is an important component of the ascorbate-glutathione cycle, which is involved in the regulation of hydrogen peroxide (H2O2) concentrations in plants. During chilling and cold acclimation, i.e. exposure to temperatures between 0 and 15 degrees C, H2O2 may accumulate. Excess electrons from the photosynthetic and respiratory electron transport chains can be used for the reduction of oxygen, thus producing superoxide radicals (O2.-); these are subsequently transformed to H2O2 via superoxide dismutase (SOD; EC 1.15.1.1). During the removal of excess H2O2, reduced glutathione (GSH) is converted to its oxidised form (GSSG), and GSH is regenerated by the activity of NADPH-dependent glutathione reductase (GR; EC 1.6.4.2). At low non-freezing temperatures, high GSH content and GR activity were detected in several plant species, indicating a possible contribution to chilling tolerance and cold acclimation. Changes in H2O2 concentration and GSH/GSSG ratio alter the redox state of the cells and may activate special defence mechanisms through a redox signalling chain. The finding that several defence genes have antioxidant responsive elements or GSSG binding sites in their regulatory regions supports the idea that redox signalling is involved in regulating gene expression in response to low temperature.
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