The tomato DELLA protein PROCERA promotes ABA responses in guard cells by upregulating ABA transporter

Abstract

Plants reduce transpiration to avoid drought stress by stomatal closure. While abscisic acid (ABA) has a central role in the regulation of stomatal closure under water-deficit conditions, we demonstrated in tomato that the gibberellin (GA) response inhibitor, the DELLA protein PROCERA (PRO), promotes ABA-induced stomatal closure and gene transcription in guard cells. To study how PRO affects stomatal closure, we performed RNAseq analysis of isolated guard cells and identified the ABA transporters ABA-IMPORTING TRANSPORTER1.1 (AIT1.1) and AIT1.2, also called in Arabidopsis NPF4.6, as upregulated by PRO. Tomato has four AIT1 genes, but only AIT1.1 and AIT1.2 were upregulated by PRO, and only AIT1.1 exhibited high expression in guard cells. Functional analysis of AIT1.1 in yeast confirmed its activity as an ABA transporter, possibly importer. CRISPR-Cas9-defrived ait1.1 mutant exhibited increased transpiration, larger stomatal aperture and reduced response to ABA. Moreover, ait1.1 suppressed the promoting effects of PRO on ABA-induced stomatal closure and gene expression in guard cells. The negative interaction between GA and ABA has been studied for many years in numerous plant species. These studies suggest that the crosstalk is mediated by changes in hormone biosynthesis and signaling. Our results suggest that it is also mediated by changes in hormone transport. One-sentence SummaryThe tomato DELLA protein PROCERA promoted abscisic acid-induced stomatal closure and gene expression by upregulating the expression of the ABA transporter ABA-IMPORTING TRANSPORTER 1 in guard cells.