Aquaporin belongs to a highly conserved group of membrane proteins called major intrinsic proteins that facilitate water transport across biological membranes. The genome of Arabidopsis encodes 35 aquaporin genes with 13 homologs in the plasma membrane intrinsic protein (PIP) subgroup. However, the function of each individual aquaporin isoform and the integrated function of plant aquaporins under various physiological conditions remain unclear. As a step toward understanding the aquaporin function in plants under various environmental stimuli, the expressions of a gene family encoding 13 PIPs in Arabidopsis thaliana under various abiotic stress conditions including drought, cold, and high salinity, or abscisic acid (ABA) treatment were investigated by a quantitative real-time reverse transcription-PCR analysis. Several PIP genes were predominantly expressed either in the roots or in the flowers. The expressions of both the highly expressed aquaporins including PIP1;1, PIP1;2, and PIP2;7 and the weakly expressed aquaporins such as PIP1;4, PIP2;1, PIP2;4, and PIP2;5 were modulated by external stimuli. The analyses of our data revealed that only the PIP2;5 was up-regulated by cold treatment, and most of the PIP genes were down-regulated by cold stress. Marked up- or down-regulation in PIP expression was observed by drought stress, whereas PIP genes were less-severely modulated by high salinity. The responsiveness of each aquaporin to ABA were different, implying that the regulation of aquaporin expression involves both ABA-dependent and ABA-independent signaling pathways. Together, our comprehensive expression profile of the 13 members of the PIP gene family provides novel basis to allocate the stress-related biological function to each PIP gene.

Although the number of reports demonstrating the roles of individual aquaporins in plants under diverse physiological conditions is expanding, the importance of interactions between different aquaporin isoforms and their integrated functions under stress conditions remain unclear. Here, we expressed one cucumber aquaporin gene, designated CsPIP1;1, and one figleaf gourd aquaporin gene, designated CfPIP2;1, in Arabidopsis thaliana, and investigated the effect of its expression on the natural expression patterns of endogenous PIP genes under stress conditions. The transcript levels of endogenous Arabidopsis PIP members were altered differently depending on stress conditions by the expression of CsPIP1;1 or CfPIP2;1. The transgenic Arabidopsis plants that constitutively express CfPIP2;1 displayed better growth compared with the wild-type plants under dehydration stress conditions, whereas CsPIP1;1 expression exerted a negative effect on the growth of Arabidopsis under dehydration stress conditions. CsPIP1;1 or CfPIP2;1 expression facilitated seed germination under high salt stress conditions, but had no influence on the growth of Arabidopsis under cold stress conditions. Our results indicate that the ectopic expression of a foreign aquaporin gene perturbs differently the natural expression patterns of endogenous aquaporin genes depending on particular stress conditions, and thereby influences the responses of plants to different stress conditions. This implies that the up- and/or down-regulation of aquaporins and their integrated functions are crucial to the maintenance of proper water balance under stress conditions.