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Systemic acquired resistance (SAR) has been reported to be associated with lesion-mimic mutants. Tobacco plants expressing vacuolar and apoplastic yeast-derived invertase (vaclnv and cwlnv, respectively) develop spontaneous necrotic lesions similar to hypersensitive responses caused by avirulent pathogens. Therefore, SAR and metabolic alterations leading to the activation of defense-related responses were studied in these plants. Defense-related gene transcripts, callose content, peroxidase activities, and levels of salicylic acid were found to be elevated. The defense reactions were accompanied by increased resistance toward potato virus Y and were measured as decreased vira1 spreading and reduced multiplication in systemic leaves of the transgenic plants. Interestingly, the accumulation of pathogenesis-related (PR) protein transcripts (PR-Q) and repression of photosynthstic gene transcripts (chlorophyll a/b binding protein) were inversely correlated and required the same threshold leve1 of hexoses for induction and repression. Expression of a cytosolic yeast-derived invertase in transgenic tobacco plants with equally increased levels of sugars neither displayed SAR responses nor showed decreased levels of photosynthetic genes. It is suggested that hexose sensing in the secretory pathway is essential for mediating the activation of defense-related genes as well as repression of photosynthetic genes in vaclnv and cwlnv plants.

Systemic acquired resistance (SAR) has been reported to be associated with lesion-mimic mutants. Tobacco plants expressing vacuolar and apoplastic yeast-derived invertase (vaclnv and cwlnv, respectively) develop spontaneous necrotic lesions similar to hypersensitive responses caused by avirulent pathogens. Therefore, SAR and metabolic alterations leading to the activation of defense-related responses were studied in these plants. Defense-related gene transcripts, callose content, peroxidase activities, and levels of salicylic acid were found to be elevated. The defense reactions were accompanied by increased resistance toward potato virus Y and were measured as decreased viral spreading and reduced multiplication in systemic leaves of the transgenic plants. Interestingly, the accumulation of pathogenesis-related (PR) protein transcripts (PR-Q) and repression of photosynthetic gene transcripts (chlorophyll a/b binding protein) were inversely correlated and required the same threshold level of hexoses for induction and repression. Expression of a cytosolic yeast-derived invertase in transgenic tobacco plants with equally increased levels of sugars neither displayed SAR responses nor showed decreased levels of photosynthetic genes. It is suggested that hexose sensing in the secretory pathway is essential for mediating the activation of defense-related genes as well as repression of photosynthetic genes in vaclnv and cwlnv plants.

No abstract

Soluble sugars have been found to regulate a number of genes involved in functions associated with sink metabolism, defense reactions and photosynthesis. As viruses and pathogens induce the expression of pathogenesis-related (PR) protein genes and have also been reported to lead to localized sugar accumulation in leaves, it was investigated whether a salicylic acid-independent but sugar-dependent pathway for PR-protein gene Induction may exist in plant cells. Leaf discs of tobacco plants were floated on different sugar solutions, transcript accumulation and salicylic acid (SA) levels were subsequently determined. PR-Q and PAR-l transcripts were found to be inducible by glucose, fructose and sucrose. No significant change in SA content could be detected, following incubation. On the other hand, SAR8.2 transcripts were repressed by elevated levels of soluble sugars and sorbitol, respectively, suggesting sensitivity to turgor pressure. Since leaves undergo sink to source transition during growth, sugar responsiveness was investigated in leaves of different developmental stages. Interestingly, induction of PR-Q and PAR-l by soluble sugars was essentially restricted to fully expanded leaves and was independent of plant age. Induction by salicylate was not confined to the source capacity of a leaf but was dependent on the age of the respective leaf. Repression of transcripts encoding photosynthetic genes (ribulose-1,5-bisphosphate carboxylaseloxygenase (r&S) and chlorophyll a/b binding protein (cub) by soluble sugars were largely independent from the leaf developmental state. These findings hint to the possibility of salicylic acid-independent defense reactions of plants against pathogens by induction of a set of PR proteins in source leaves. Furthermore, the data suggest different mechanisms for the induction of PR-protein genes and the repression of photosynthetic genes by soluble sugars.

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Roots of intact 5-day-old maize (Zea mays L.) seedlings were exposed to 3 micromolar Cd during a 7-day period. Cysteine, gamma-glutamylcysteine, glutathione (GSH), and Cd-induced acid-soluble thiols (ASTs), including phytochelatins, were quantified in roots and shoots. Adaptation to Cd and its cost to seedling development were evaluated by measuring Cd content, tissue fresh weight, and rate of root elongation. Roots contained 60 to 67% of the Cd in the seedlings between 4 and 7 days of exposure. Exposure to Cd decreased the fresh weight gain in roots from day 4 onward without affecting the shoots. Between days 1.5 and 3.5 of Cd treatment, roots elongated more slowly than controls; however, their growth rate recovered thereafter and exceeded that of controls. Exposure to Cd did not appreciably affect the concentration of cysteine in the seedlings. However, the initial low concentration of gamma-glutamylcysteine increased (after a lag of 6 hours in roots and 2 days in shoots), reaching a plateau by day 6 at 28.5 nanomoles per gram of fresh weight in roots and by day 5 at 19.1 nanomoles per gram of fresh weight in shoots. During the first 9 hours of Cd exposure, the concentration of GSH in roots decreased dramatically (at 31.6 nanomoles per gram of fresh weight per hour) and thereafter decreased more slowly than in controls. The depletion of GSH in the roots (366 nanomoles per gram of fresh weight) matched the synthesis of ASTs (349 nanomoles per gram of fresh weight) during the first 48 hours. The concentration of ASTs in roots increased steadily thereafter to reach 662.2 nanomoles per gram of fresh weight by 6 days of Cd exposure. In shoots, Cd had little influence on the concentration of GSH, but ASTs still accumulated to 173.3 nanomoles per gram fresh weight after 5 days. The molar ratio of thiols in ASTs to Cd increased to a maximum of 10.24 in roots after 4 hours and of 4.25 in shoots after 2 days of Cd exposure. After 4 days, the ratio reached a plateau of approximately 2 in roots and between 2 and 3 in shoots, as if a steady state of Cd chelation had been achieved in both organs. The plateau coincided with recovered root elongation or an adaptation to Cd. The reduced fresh weight gain of the roots during this time, however, indicated that the synthesis of Cd-induced thiols was at a cost to root development.