GNOM-Mediated Vesicular Trafficking Plays an Essential Role in Hydrotropism of Arabidopsis Roots

Roots respond not only to gravity but also to moisture gradient by displaying gravitropism and hydrotropism, respectively, to control their growth orientation, which helps plants obtain water and become established in the terrestrial environment. As gravitropism often interferes with hydrotropism, however, the mechanisms of how roots display hydrotropism and differentiate it from gravitropism are not understood. We previously reported MIZU-KUSSEI1 (MIZ1) as a gene required for hydrotropism but not for gravitropism, although the function of its protein was not known. Here, we found that a mutation of GNOM encoding guanine-nucleotide exchange factor for ADP-ribosylation factor-type G proteins was responsible for the ahydrotropism of Arabidopsis (Arabidopsis thaliana), miz2. Unlike other gnom alleles, miz2 showed no apparent morphological defects or reduced gravitropism. Instead, brefeldin A (BFA) treatment inhibited both hydrotropism and gravitropism in Arabidopsis roots. In addition, a BFA-resistant GNOM variant, GN^M696L, showed normal hydrotropic response in the presence of BFA. Furthermore, a weak gnom allele, gnom^B/E, showed defect in hydrotropic response. These results indicate that GNOM-mediated vesicular trafficking plays an essential role in hydrotropism of seedling roots.Stationary growth is a distinct feature of plants and distinguishes them from other organisms. Plants have evolved a variety of mechanisms for responding to environmental cues, which enables them to survive in the presence of limited resources or environmental stresses. One of the most important growth adaptations plants have acquired is tropism, growth response that involves bending or curving of plant organs toward or away from a stimulus. For example, roots display tropisms in response to environmental cues such as gravity, light, touch, and moisture (Darwin and Darwin, 1880; Takahashi, 1997; Correll and Kiss, 2002; Monshausen et al., 2008). Gravitropism has been the subject of intense study, while other tropic responses of roots have been less well characterized. There is some evidence of hydrotropism in roots, but this response has proven difficult to differentiate from gravitropism, as the latter always interferes with hydrotropism (Jaffe et al., 1985; Takahashi, 1994; Takahashi, 1997). The demonstration of true hydrotropism in roots has facilitated the identification of some of the physiological aspects of hydrotropism and its existence in a wide range of plant species. However, the underlying mechanisms that regulate hydrotropism remain unknown. The limited supply of water and precipitation in many parts of the world greatly affects agriculture and ecosystems. Elucidating the molecular mechanism of hydrotropism in roots is therefore important not only for understanding how terrestrial plants adapt to changes in moisture, but also for improving crop yields and biomass production.The isolation and analysis of hydrotropism-deficient mutants using the model plant species Arabidopsis (Arabidopsis thaliana) represents a potent tool for dissecting the molecular mechanism of hydrotropism. Previously, we isolated an ahydrotropic mutant of Arabidopsis, mizu-kussei1 (miz1), and showed that MIZ1 encodes a protein of unknown function (Kobayashi et al., 2007). In light of both the physiological features of hydrotropism, as well as what we have learned from genetic studies of other tropisms, it is unlikely that miz1 alone governs the hydrotropic response. In support of this, we have identified a second ahydrotropic mutant, miz2, a unique allele of gnom that confers ahydrotropic but not agravitropic growth, which implies distinct roles of vesicular trafficking between hydrotropism and gravitropism in roots.