The tomato Cu,Zn superoxide dismutase genes are developmentally regulated and respond to light and stress

The expression of the two Cu,Zn superoxide dismutase (SOD) genes of tomato was followed in different organs and plant developmental stages at the transcript and enzymatic activity levels. The cDNA clones used as probes code for the chloroplast Cu,Zn SOD (clone T1) and the cytosolic Cu,Zn SOD (clone P31). The two genes were found to display distinct expression patterns. While the T1 transcript was rare or absent from roots, stems and ripening fruits, the P31 transcript was very abundant in these organs. Shoot tips, flower buds, seedlings and young leaves contained high levels of the two mRNAs. During leaf expansion, the levels of both transcripts diminish markedly. Despite the diminished presence of transcripts, SOD activity levels of the corresponding cytosolic and chloroplast isozymes accumulated and were sustained throughout leaf expansion. In non-photosynthetic organs, the SOD-3 (cytosolic) isozyme contained most of the activity, while in the expanded leaf the SOD-1 (chloroplast) isozyme was more abundant. Light-regulated accumulation of both the P31 transcript (1.7-fold) and the T1 transcript (3-fold) was observed upon light exposure of etiolated seedlings. However, only SOD-1 activity was observed to increase, after a lag of a few hours. The levels of both transcripts increased in response to paraquat and mechanical wounding. The level of the cytosolic transcript and the respective isozyme activity increased dramatically during prolonged drought stress while the chloroplast transcript remained unaffected. The expression of both genes was enhanced by spraying tomato plants with ethephon--a compound that releases ethylene. Our data show that the expression of Cu,Zn SOD genes in tomato is modulated in response to a variety of factors and suggest the importance of oxyradical toxicity as well as the role of SOD in the defence mechanism of plants exposed to stress.