Transcriptional regulation by abscisic acid in barley (Hordeum vulgare L.) seeds involves autoregulation of the transcription factor HvABI5

The barley bZIP transcription factor HvABI5 mediates abscisic acid (ABA)-upregulated gene expression in barley (Hordeum vulgare L.) seeds. HvABI5 specifically recognizes cis-elements of the ABA response complexes present in the promoters of the ABA-induced genes HVA1 and HVA22. HvABI5 together with another transcription factor, HvVP1, are required for the transactivation of these promoters, and this transactivation process is insensitive to the negative regulator abi1-1. The expression of HvABI5 itself appeared to be induced by ABA and can be suppressed by abi1-1. Gain- and loss-of-function studies in barley aleurone cells show that HvABI5 expression is positively regulated by a feed-forward circuit that involves HvABI5 itself and HvVP1. Mutation of the Ser residue in HvABI5, which has been shown to be phosphorylated in an ABA-dependent manner in the rice orthologue of HvABI5, reduces the transactivation activity of the factor by 50%. Although levels of HvABI5 and its transcript are enhanced by ABA treatment, the nuclear localization of HvABI5 is not affected by ABA. A model based on these observations is presented to explain the ABA upregulation of gene expression.