Regulation of Sulfur Nutrition in Wild-Type and Transgenic Poplar Over-Expressing γ-Glutamylcysteine Synthetase in the Cytosol as Affected by Atmospheric H(2)S

This study with poplar (Populus tremula × Populus alba) cuttings was aimed to test the hypothesis that sulfate uptake is regulated by demand-driven control and that this regulation is mediated by phloem-transported glutathione as a shoot-to-root signal. Therefore, sulfur nutrition was investigated at (a) enhanced sulfate demand in transgenic poplar over-expressing γ-glutamylcysteine (γ-EC) synthetase in the cytosol and (b) reduced sulfate demand during short-term exposure to H(2)S. H(2)S taken up by the leaves increased cysteine, γ-EC, and glutathione concentrations in leaves, xylem sap, phloem exudate, and roots, both in wild-type and transgenic poplar. The observed reduced xylem loading of sulfate after H(2)S exposure of wild-type poplar could well be explained by a higher glutathione concentration in the phloem. In transgenic poplar increased concentrations of glutathione and γ-EC were found not only in leaves, xylem sap, and roots but also in phloem exudate irrespective of H(2)S exposure. Despite enhanced phloem allocation of glutathione and its accumulation in the roots, sulfate uptake was strongly enhanced. This finding is contradictory to the hypothesis that glutathione allocated in the phloem reduces sulfate uptake and its transport to the shoot. Correlation analysis provided circumstantial evidence that the sulfate to glutathione ratio in the phloem may control sulfate uptake and loading into the xylem, both when the sulfate demand of the shoot is increased and when it is reduced.