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The role of the redox state of the apoplast in hormone responses, signaling cascades, and gene expression was studied in transgenic tobacco (Nicotiana tabacum) plants with modified cell wall-localized ascorbate oxidase (AO). High AO activity specifically decreased the ascorbic acid (AA) content of the apoplast and altered plant growth responses triggered by hormones. Auxin stimulated shoot growth only when the apoplastic AA pool was reduced in wild-type or AO antisense lines. Oxidation of apoplastic AA in AO sense lines was associated with loss of the auxin response, higher mitogen-activated protein kinase activities, and susceptibility to a virulent strain of the pathogen Pseudomonas syringae. The total leaf glutathione pool, the ratio of reduced glutathione to glutathione disulfide, and glutathione reductase activities were similar in the leaves of all lines. However, AO sense leaves exhibited significantly lower dehydroascorbate reductase and ascorbate peroxidase activities than wild-type and antisense leaves. The abundance of mRNAs encoding antioxidant enzymes was similar in all lines. However, the day/night rhythms in the abundance of transcripts encoding the three catalase isoforms were changed in response to the AA content of the apoplast. Other transcripts influenced by AO included photorespiratory genes and a plasma membrane Ca 21 channel-associated gene. We conclude that the redox state of the apoplast modulates plant growth and defense responses by regulating signal transduction cascades and gene expression patterns. Hence, AO activity, which modulates the redox state of the apoplastic AA pool, strongly influences the responses of plant cells to external and internal stimuli.

Plants have evolved a plethora of mechanisms to circumvent the potential damaging effects of living under low phosphorus availability in the soil. These mechanisms include different levels of organization, from root-shoot signalling at the whole-plant level to specific biochemical responses at the subcellular level, such as reductions in photosynthesis and the consequent activation of photo- and antioxidant mechanisms in chloroplasts. Some recent studies clearly indicate that severe phosphorus deficiency can lead to alterations in the photosynthetic apparatus, including reductions in CO2 assimilation rates, a down-regulation of photosynthesis-related genes and photoinhibition at the photosystem II level, thus causing potential photo-oxidative stress. Photo-oxidative stress is characterized by an increased production of reactive oxygen species in chloroplasts, which at low concentrations can serve a signalling, protective role, but when present at high concentrations can cause damage to lipids, proteins and nucleic acids, thus leading to irreversible injuries. We discuss here the mechanisms that phosphate-starved plants have evolved to withstand photo-oxidative stress, including changes at the subcellular level (e.g. activation of photo- and antioxidant protection mechanisms in chloroplasts), cellular and tissular levels (e.g. activation of photorespiration and anthocyanin accumulation) and whole-plant level (alterations in source-sink relationships modulated by hormones). Of particular importance is the current evidence demonstrating that phosphate-starved plants activate simultaneous responses at multiple levels, from transcriptional changes to root-shoot signalling, to prevent oxidative damage. In this review, we summarize current knowledge about the occurrence of photo-oxidative stress in phosphate-starved plants and highlight the mechanisms these plants have evolved to prevent oxidative damage under phosphorus limitation at the subcellular, cellular and whole-plant levels.

Mediterranean plants have evolved a complex antioxidant defense system to cope with summer drought. Flavonoids, and particularly flavanols and flavonols, are potent in vitro antioxidants, but their in vivo significance within the complex network of antioxidant defenses remains unclear, especially in plant responses to stress. To gain insight into the role of flavonoids in the antioxidant defense system of Cistus clusii Dunal, we evaluated drought-induced changes in flavonoids in leaves and compared the response of these compounds with that of other low molecular weight antioxidants (ascorbic acid, tocopherols and carotenoids). Among the antioxidant flavonoids analyzed, epigallocatechin gallate was present in the greatest concentrations (up to about 5 micromol dm(-2)). Other flavanols, such as epicatechin and epicatechin gallate, were found at concentrations below 0.25 and 0.03 micromol dm(-2), respectively. Neither of the antioxidant flavonols analyzed, quercetin and kaempferol, were detected in C. clusii leaves. Epigallocatechin gallate, ascorbic acid and alpha-tocopherol concentrations increased to a similar extent (up to 2.8-, 2.6- and 3.3-fold, respectively) in response to drought, but the kinetics of the drought-induced increases differed. Epigallocatechin gallate, epicatechin and epicatechin gallate concentrations increased progressively during drought, reaching maximum values after 30 days of stress. Ascorbic acid concentrations increased twofold after 15 days of drought, and maximum values were attained after 50 days of drought. In contrast, alpha-tocopherol concentrations remained constant during the first 30 days of drought, but increased sharply by 3.3-fold after 50 days of drought. The maximum efficiency of photosystem II photochemistry and the extent of lipid peroxidation remained constant throughout the drought period, whereas the redox state of ascorbic acid and alpha-tocopherol shifted toward their reduced forms in response to drought, indicating that the concerted action of low molecular weight antioxidants may help prevent oxidative damage in plants.

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Salt stress is one of the most important abiotic stress factors affecting plant growth and productivity in natural ecosystems. In this study, we aimed at determining possible differences between salt tolerant and salt sensitive species in early (within 72 h) salt stress response in leaves and roots. To this purpose, we subjected three Brassicaceae species, namely two halophytes-Cakile maritima and Thellungiella salsuginea--and a glycophyte--Arabidopsis thaliana- to short-term salt stress (400 mM NaCl). The results indicate that the halophytes showed a differential osmotic and ionic response together with an early and transient oxidative burst, which was characterized by enhanced hydrogen peroxide levels and subsequent activation of antioxidant defenses in both leaves and roots. In addition, the halophytes displayed enhanced accumulation of abscisic acid, jasmonic acid (JA) and ACC (aminocyclopropane-1-carboxylic acid, the precursor of ethylene) in leaves and roots, as compared to A. thaliana under salt stress. Moreover, the halophytes showed enhanced expression of ethylene response factor1 (ERF1), the convergence node of the JA and ethylene signaling pathways in both leaves and roots upon exposure to salt stress. In conclusion, we show that the halophytes C. maritima and T. salsuginea experience an early oxidative burst, improved antioxidant defenses and hormonal response not only in leaves but also in roots, in comparison to the glycophyte A. thaliana. This differential signaling response converging, at least in part, into increased ERF1 expression in both above- and underground tissues seems to underlay, at least in part, the enhanced tolerance of the two studied halophytes to salt stress.

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