The oxidation of ABI4 by RBOHD-derived reactive oxygen species integrates redox signaling into abscisic-acid and drought-stress responses

The phytohormone abscisic acid (ABA) induces stomatal closure and facilitates plant adaptation to drought stress. Reactive oxygen species (ROS) produced by the NADPH oxidase RESPIRATORY BURST OXIDASE HOMOLOG PROTEIN D (RBOHD) are essential components of ABA signal transduction. However, the molecular mechanisms by which ABA influences ROS production, and how ROS signaling is integrated into the ABA signaling network, are poorly understood. Here, we report that the transcription factor ABSCISIC ACID INSENSITIVE 4 (ABI4) acts upstream of RBOHD to regulate ABA-induced ROS production and stomatal closure in Arabidopsis ( Arabidopsis thaliana ). We determined that ABA-induced RBOHD expression and ROS production were significantly attenuated in the abi4 mutant. In the wild type, an ABA-triggered ROS burst promoted the sulfenylation of ABI4 at Cys250, which enhanced its DNA binding and transactivation activity toward RBOHD , thereby regulating ABA signaling. The sulfenylation-dependent regulation of ABI4 was required for ABA-induced stomatal closure and greatly contributed to plant drought tolerance. These findings uncover a redox-dependent feedback loop in which ABI4 oxidation fine-tunes ABA responses and drought tolerance by dynamically regulating RBOHD-mediated ROS production. This post-translational regulatory mechanism for ABA signaling establishes a molecular framework explaining the crosstalk between ABA and ROS in the regulation of stomatal closure and drought resilience.