myo-Inositol and its derivatives are commonly studied with respect to cell signaling and membrane biogenesis, but they also participate in responses to salinity in animals and plants. In this study, we focused on L -myo -inositol 1-phosphate synthase (INPS), which commits carbon to de novo synthesis, and myo -inositol O -methyltransferase (IMT), which uses myo -inositol for stress-induced accumulation of a methylinositol, D -ononitol. The Imt and Inps promoters are transcriptionally controlled. We determined that the transcription rates, transcript levels, and protein abundance are correlated. During normal growth, INPS is present in all cells, but IMT is repressed. After salinity stress, the amount of INPS was enhanced in leaves but repressed in roots. IMT was induced in all cell types. The absence of myo -inositol synthesis in roots is compensated by inositol/ononitol transport in the phloem. The mobilization of photosynthate through myo -inositol translocation links root metabolism to photosynthesis. Our model integrates the transcriptional control of a specialized metabolic pathway with physiological reactions in different tissues. The tissue-specific differential regulation of INPS, which leads to a gradient of myo -inositol synthesis, supports root growth and sodium uptake. By inducing expression of IMT and increasing myo -inositol synthesis, metabolic end products accumulate, facilitating sodium sequestration and protecting photosynthesis.
INTRODUCTIONA central focus of our work is to determine genes that underlie mechanisms providing salinity tolerance in higher plants. In addition, we hope to learn how whole plants adapt metabolically. Many researchers have studied tolerance mechanisms in plants. Their findings show the importance of metabolite accumulation, which generally is attributed to osmotic adjustment leading to water retention and/or protection of biochemical pathways. When single enzymes were tested in transgenic plants, the accumulation of mannitol, proline, fructans, trehalose, glycine betaine, or ononitol (Tarczynski et al., 1993;Kavi Kishor et al., 1995;Pilon-Smits et al., 1995;Holmström et al., 1996;Hayashi et al., 1997;Sheveleva et al., 1997) provided marginally higher salinity and cold or drought tolerance. The manipulation of biochemical pathways by the transfer of multiple genes will be an even more appropriate strategy that is expected to provide broader tolerance because salinity tolerance is a multigenic trait Warne et al., 1995;Bohnert and Jensen, 1996). Salinity tolerance is also a multicellular trait, and the remodeling of complex pathways in transgenic plants will require knowledge of the distribution of such pathways throughout the plant. We have analyzed the myo -inositol biosynthetic pathway-leading to methylated inositols-that feeds into a pathway specific to salinity tolerance in a halophyte, the common ice plant ( Mesembryanthemum crystallinum ) (Loewus and Dickinson, 1982;Vernon and Bohnert, 1992).myo -Inositol is a central component of several biochemical pathways. It i...