ZmMKK4, a novel group C mitogen-activated protein kinase kinase in maize (Zea mays), confers salt and cold tolerance in transgenic Arabidopsis

Mitogen-activated protein kinase (MAPK) cascades are signalling modules that transduce extracellular signalling to a range of cellular responses. Plant MAPK cascades have been implicated in development and stress response. In this study, we isolated a novel group C MAPKK gene, ZmMKK4, from maize. Northern blotting analysis revealed that the ZmMKK4 transcript expression was up-regulated by cold, high salt and exogenous H2O2, but down-regulated by exogenous abscisic acid (ABA). Over-expression of ZmMKK4 in Arabidopsis conferred tolerance to cold and salt stresses by increased germination rate, lateral root numbers, plant survival rate, chlorophyll, proline and soluble sugar contents, and antioxidant enzyme [peroxidase (POD), catalase (CAT)] activities compared with control plants. Furthermore, ZmMKK4 enhanced a 37 kDa kinase activity after cold and salt stresses. RT-PCR analysis revealed that the transcript levels of stress-responsive transcription factors and functional genes were higher in ZmMKK4-over-expressing plants than in control plants. In addition, ZmMKK4 protein is localized in the nucleus. Taken together, these results indicate that ZmMKK4 is a positive regulator of salt and cold tolerance in plants.