Tonoplast Intrinsic Proteins (TIP) are one of five subfamilies of aquaporins in higher plants. Plants typically contain a large number of TIP genes, ranging from 6 to 35 compared to humans. The molecular weight of the TIP subfamily members ranges from 25 to 28 kDa. Despite their sequence diversity, all TIP monomers have the same structure, which consists of six transmembrane helices and five inter-helical loops that form an hourglass shape with a central pore. Four monomers form tetramers, which are functional units in the membrane. TIPs form channels in the tonoplast that basically function as regulators of the intracellular water flow, which implies that they have a role in regulating cell turgor. TIPs are responsible for precisely regulating the movement of not only water, but also some small neutral molecules such as glycerol, urea, ammonia, hydrogen peroxide and formamide. The expression of TIPs may be affected by different environmental stresses, including drought, salinity and cold. TIPs expression is also altered by phytohormones and the appropriate cis-regulatory motifs are identified in the promotor region of the genes encoding TIPs in different plant species. It was shown that manipulating TIP-encoding genes expression in plants could have the potential to improve abiotic stress tolerance.
Tobacco chloroplast transformants expressing genes encoding dehydroascorbate reductase, glutathione reductase, and glutathione-S-transferase, exhibit altered anti-oxidant metabolism and improved abiotic stress tolerance
