The Arabidopsis transcription factor ABSCISIC ACID INSENSITIVE 4 (ABI4) is a key player in the plant hormone abscisic acid (ABA) signaling pathway and is involved in plant response to abiotic stress and development. Expression of the ABI4 gene is tightly regulated, with low basal expression. Maximal transcript levels occur during the seed maturation and early seed germination stages. Moreover, ABI4 is an unstable, lowly expressed protein. Here, we studied factors affecting the stability of the ABI4 protein using transgenic Arabidopsis plants expressing 35S::HA-FLAG-ABI4-eGFP. Despite the expression of eGFP-tagged ABI4 being driven by the highly active 35S CaMV promoter, low steady-state levels of ABI4 were detected in the roots of seedlings grown under optimal conditions. These levels were markedly enhanced upon exposure of the seedlings to abiotic stress and ABA. ABI4 is degraded rapidly by the 26S proteasome, and we report on the role of phosphorylation of ABI4-serine 114 in regulating ABI4 stability. Our results indicate that ABI4 is tightly regulated both post-transcriptionally and post-translationally. Moreover, abiotic factors and plant hormones have similar effects on ABI4 transcripts and ABI4 protein levels. This double-check mechanism for controlling ABI4 reflects its central role in plant development and cellular metabolism.
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The transcription factor ABA-INSENSITIVE(ABI)4 has diverse roles in regulating plant growth, including inhibiting germination and reserve mobilization in response to ABA and high salinity, inhibiting seedling growth in response to high sugars, inhibiting lateral root growth, and repressing light-induced gene expression. ABI4 activity is regulated at multiple levels, including gene expression, protein stability, and activation by phosphorylation. Although ABI4 can be phosphorylated at multiple residues by MAPKs, we found that S114 is the preferred site of MPK3. To examine the possible biological role of S114 phosphorylation, we transformed abi4-1 mutant plants with ABI4pro::ABI4 constructs encoding wild type (114S), phosphorylation-null (S114A) or phosphomimetic (S114E) forms of ABI4. Phosphorylation of S114 is both necessary and sufficient for the response to ABA, glucose, salt stress, and lateral root development, where the abi4 phenotype could be complemented by expressing ABI4(114S) or ABI4(S114E) but not ABI4(S114A). Comparison of root transcriptomes in ABA-treated roots of abi4-1 mutant plants transformed with constructs encoding the different phosphorylation-forms of S114 of ABI4 revealed that 85% of the ABI4-regulated genes whose expression pattern could be restored by expressing ABI4(114S) are down-regulated by ABI4. Over half of the ABI4-modulated genes were independent of the phosphorylation state of ABI4; these are enriched for stress responses. Phosphorylation of S114 was required for regulation of 35% of repressed genes, but only 17% of induced genes. The genes whose repression requires the phosphorylation of S114 are mainly involved in embryo and seedling development, growth and differentiation, and regulation of gene expression.
The Arabidopsis transcription factor ABSCISIC ACID INSENSITIVE 4 (ABI4) is a key player in the plant hormone abscisic acid (ABA) signaling pathway. ABI4 is also involved in seed development and germination, the response to abiotic stresses such as drought and salinity, control of lipid reserve mobilization in the embryo, lateral root formation, and redox control. Expression of the ABI4 gene is tightly regulated and basal expression is low. Maximal transcript levels occur during seed maturation and in the early stages of seed germination and are markedly reduced in other developmental stages. ABI4 is an unstable lowly expressed protein, resulting from tight post-transcriptional regulation. Here, we studied factors affecting the stability of the ABI4 protein using transgenic Arabidopsis plants expressing 35S::HA-FLAG-ABI4-eGFP. Despite the expression of eGFP-tagged ABI4 being driven by the highly active 35S CaMV promoter the steady-state levels of ABI4 were extremely low in the roots of seedling grown in optimal conditions. These levels were markedly enhanced upon exposure of the seedlings to abiotic stress and ABA. ABI4 is degraded rapidly by the 26S proteasome and we report on the role of phosphorylation of ABI4-serine 114 in regulating ABI4 stability. Our results indicate that ABI4 is tightly regulated both post-transcriptionally and post-translationally. Moreover, abiotic factors and plant hormones have similar effects on ABI4 transcripts and ABI4 protein levels. This double-check mechanism for controlling ABI4 reflects on its central role in plant development and cellular metabolism.
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