Model of GCN1-GCN2-p-eIF2α in cold response. Upon cold stress or nutrient deprivation, at least three cellular signalling pathways, including those involving the ratio of AMP/ATP, GCN1-GCN2-p-eIF2 and TOR kinase, contribute to the inhibition of protein translation in Arabidopsis. In this study, GCN1-GCN2 and the TOR pathways, indicated by bold characters, were found to be required for cold tolerance. Dotted arrows indicate the pathways that are known to be involved in the cold tolerance in mammals, but the functions of these pathways in the cold tolerance in plants are still unknown. The functions of GCN1-GCN2-p-eIF2α TOR pathways in response to nutrient deficiency, such as amino acid or glucose starvation, have been well documented in Arabidopsis.