Inhibition of gibberellin accumulation by water deficiency promotes fast and long-term 'drought avoidance' responses in tomato

Plants reduce transpiration to avoid dehydration during drought episodes by stomatal closure and inhibition of canopy growth. While abscisic acid (ABA) has a primary role in drought avoidance, previous studies suggest that gibberellin (GA), might also be involved. Here we show in tomato (Solanum lycopersicum) that shortage of water inhibited the expression of the GA biosynthesis genes GA20 oxidase1 (GA20ox1) and GA20ox2 and induced the GA-deactivating gene GA2ox7 in leaves and guard cells, resulting in reduced bioactive GA levels. Drought regulation of GA metabolism was mediated by ABA-dependent and independent pathways, and by the transcription factor DEHYDRATION RESPONSIVE ELEMENT BINDING (DREB), TINY1. Mutations in GA20ox1 and GA20ox2 reduced water loss due to the smaller canopy area. On the other hand, loss of GA2ox7 did not affect leaf size, but attenuated stomatal response to water deficiency; during soil dehydration, ga2ox7 plants closed their stomata and reduced transpiration later than WT, suggesting that ga2ox7 stomata are hyposensitive to soil dehydration. Together, the results suggest that drought-induced GA deactivation in guard cells contributes to stomatal closure at the early stages of soil dehydration, whereas inhibition of GA synthesis in leaves promotes mainly the long-term reduction in canopy growth to reduce transpiration area.