In many plants, free proline accumulates in response to the imposition of a wide range of biotic and abiotic stresses. Controversy has surrounded the extent to which this shift in nitrogen metabolism benefits plants under adverse environmental conditions. Most attempts to account for the phenomenon have focused on the ability of proline to mediate osmotic adjustment, stabilise subcellular structures and scavenge free radicals. However, often the cytoplasmic pool of free proline even after the imposition of stress is insufficient size to account for pronounced biophysical effects. Alternatively, selective preservation of this stress-induced response may relate to endpoints other than simply augmenting the cellular pool of free proline. Proline accumulation may reduce stress-induced cellular acidification or prime oxidative respiration to provide energy needed for recovery. High levels of proline synthesis during stress may maintain NAD(P)+/NAD(P)H ratios at values compatible with metabolism under normal conditions. Consideration of the cofactor preference of plant Δ1-pyrroline-5-carboxylate (P5C) reductase as well as the in vivo concentrations of the two pyridine nucleotide cofactors and their respective redox ratios suggests that even a small increase in proline biosynthesis might have a large impact on the level of reduction of the cellular NADP pool. The increased NADP+/NADPH ratio mediated by proline biosynthesis is likely to enhance activity of the oxidative pentose phosphate pathway. This would provide precursors to support the demand for increased secondary metabolite production during stress as well as nucleotide synthesis accompanying the accelerated rate of cell division upon relief from stress, when oxidation of proline is likely to provide an important energy source for ADP phosphorylation. Thus, the extreme sensitivity of the metabolic processes of proline synthesis and degradation themselves may be of benefit by regulating metabolic processes adversely affected by stress. This viewpoint is supported by consideration of other physiological phenomena not directly related to stress responses, but in which proline metabolism may also play a regulatory role. A mechanism is proposed whereby the interconversions of proline and P5C in different cell types and the associated transfer of redox potential between tissues may constitute a form of metabolic signalling within higher plants. Stress-related alterations in proline metabolism may impinge on systems of redox control of plant gene expression.

Metabolic implications of stress-induced proline accumulation in plants

Plant cell walls are amazingly complex amalgams of carbohy? drates, proteins, lignin, water, and incrusting substances such as cutin, suberin, and certain inorganic compounds that vary among plant species, cell types, and even neighboring cells Developmental events and exposure to any of a number of abiotic and biotic stresses further increase this compositional and structural variation Moreover, the dynamic nature and func? tions of plant cell walls in terms of growth and development, environmental sensing and signaling, plant defense, intercel? lular communication, and selective exchange interfaces are reflected in these variations Much is currently known about the structure and metabolic regulation of the various cell wall components, but relatively little is known about their precise functions and intermolecular interactions In this review, I will discuss the accumulated structural and regulatory data and the much more limited functional and in? termolecular interaction information on five plant cell wall protein classes These five protein classes, listed in Table 1, include the extensins, the glycine-rich proteins (GRPs), the proline-rich proteins (PRPs), the solanaceous lectins, and the arabinogalactan proteins (AGPs) These five proteins may be evolutionarily related to one another, most obviously because each of them, with the exception of the GRPs, contains hydroxyproline, and less obviously in the case of the GRPs because this class has nucleotide sequence similarity to the extensins For completeness, I should mention that these are not the only cell wall proteins that are known Others exist, such as cysteine-rich thionins, 28and 70-kD water-regulated proteins, a histidine-tryptophan-rich protein, and many cell wall enzymes such as peroxidases, phosphatases, invertases, a-mannosidases, P-mannosidases, p-1,3-glucanases, (3-1,4glucanases, polygalacturonase, pectin methylesterases, malate dehydrogenase, arabinosidases, a-galactosidases, (3-galactosidases, |3-glucuronosidases, p-xylosidases, proteases, and ascorbic acid oxidase (Varner and Lin, 1989) However, the above five classes generally represent the most abundant, and to date, the most well-studied and widely documented, plant cell wall proteins Before describing these five wall protein classes, I should point out that research on these individual proteins has oc? curred in several plant species, but relatively few examples exist where these cell wall proteins have been studied together in one plant, let alone in one particular plant organ or type of cell Thus, data from one plant species are often extrapolated to represent the situation in other plant species Although such extrapolations are usually valid, enough variations are now known that caution should be exercised in making or believing such claims

Structure and function of plant cell wall proteins.

https://www.cell.com/trends/biochemical-sciences/pdf/S0968-0004(11)00201-5.pdf

How do plants feel the heat

Nitric oxide (NO) is a very active molecule involved in many and diverse biological pathways where it has proved to be protective against damages provoked by oxidative stress conditions. In this work, we studied the effect of two NO donors, sodium nitroprusside (SNP) and S-nitroso-N-acetylpenicillamine SNP-treated on the response of wheat (Triticum aestivum) to water stress conditions. After 2 and 3 h of drought, detached wheat leaves pretreated with 150 microM SNP retained up to 15% more water than those pretreated with water or NO(2)(-)/NO(3)(-). The effect of SNP treatment on water retention was also found in wheat seedlings after 7 d of drought. These results were consistent with a 20% decrease in the transpiration rate of SNP-treated detached wheat leaves for the same analyzed time. In parallel experiments, NO was also able to induce a 35%, 30%, and 65% of stomatal closure in three different species, Tradescantia sp. (monocotyledonous) and two dicotyledonous, Salpichroa organifolia and fava bean (Vicia faba), respectively. In SNP-treated leaves of Tradescantia sp., the stomatal closure was correlated with a 10% increase on RWC. Ion leakage, a cell injury index, was 25% lower in SNP-treated wheat leaves compared with control ones after the recovery period. Carboxy-PTIO (2-(4-carboxyphenyl)-4,4,5,5-tetramethylimidazoline-1-oxyl-3-oxide), a specific NO scavenger, reverted SNP action by restoring the transpiration rate, stomatal aperture, and the ion leakage to the level found in untreated leaves. Northern-blot analysis showed that SNP-treated wheat leaves display a 2-fold accumulation of a group three late embryogenesis abundant transcript with respect to control leaves both after 2 and 4 h of drought periods. All together, these results suggest that the exogenous application of NO donors might confer an increased tolerance to severe drought stress conditions in plants.

/pdf/nitric-oxide-induces-stomatal-closure-and-enhances-the-34gomqxjzz.pdf

Nitric Oxide Induces Stomatal Closure and Enhances the Adaptive Plant Responses against Drought Stress

The aim of this review is to present the current state of knowledge on plant chitinases and their regulation and function. Chitinases are up-regulated by a variety of stress conditions, both biotic and abiotic, and by such phytohormones as ethylene, jasmonic acid, and salicylic acid. Like other PR proteins, chitinases play a role in plant resistance against distinct pathogens. Moreover, by reducing the defence reaction of the plant, chitinases allow symbiotic interaction with nitrogen-fixing bacteria or mycorrhizal fungi. However, recent investigations have shown that these enzymes are also involved in numerous physiological events. The involvement of chitinases in development and growth processes is also described.

Plant chitinases--regulation and function.

Plants respond to pathogen infection and environmental stress by regulating the coordinate expression of many stress-related genes. In plants, the expression of the glycolytic enzyme glyceraldehyde-3-phosphate dehydrogenase (GAPDH) is induced under environmental stress. This work was aimed at investigating whether the expression pattern of cytosolic GAPDH is also modulated upon infection of potato plants (Solanum tuberosum L.) with the late blight fungal agent Phytophthora infestans. Northern blot analysis showed the accumulation of the GAPDH gene transcripts in leaves and stems of inoculated potato plants. When tuber discs were treated with eicosapentaenoic acid (EPA), an elicitor found in P. infestans, GAPDH gene transcripts level increased. This increase was parallel to that of the hydroxymethyl glutharyl coenzyme A reductase (HMGR), an enzyme involved in pathogen defense reactions. Glucans obtained from P. infestans cell wall acts sinergistically with EPA on GAPDH and HMGR gene induction. Salicylic acid, an endogenous signal for inducing systemic acquired resistance, was also effective in stimulating the GAPDH transcript accumulation in potato leaves. These experiments suggest that related multi-component factors, which are part of both primary and secondary metabolism, are probably regulated by similar signal transduction pathways when they are induced under biotic or abiotic stress conditions.

Accumulation of cytosolic glyceraldehyde-3-phosphate dehydrogenase RNA under biological stress conditions and elicitor treatments in potato.

The mechanism involved in systemic acquired resistance (SAR) can be non-specifically induced in susceptible plants. In response to pathogens, plants' natural defence mechanisms include the production of lignin and phytoalexins and the induction of plant enzymes. The aim of this research was to study the induction of SAR mediated by the chemical activator DL-3-aminobutyric acid (BABA) and the fungicide fosetyl-aluminium in potato cultivars with different levels of resistance against Phytophthora infestans (Mont) de Bary. To study the chemical induction of the resistance, the foliage of several potato cultivars was sprayed with BABA, fosetyl-aluminium or water (as a control treatment). After 3 days the foliage was inoculated with P. infestans. Seven days after inoculation, development of disease symptoms in the foliage was assessed. In postharvest tuber samples, evidence for enhancement of the defence response was evaluated by measuring the protein content of several hydrolytic enzymes as well as the phenol and phytoalexin content. The highest level of protection against late blight was observed when the chemicals were applied at early stages of crop development. An increase in resistance to late blight was also detected in tubers after harvest. There was also an increase in the protein level of beta-1,3-glucanase and aspartic protease as well as in the phenol and phytoalexin content of potato tuber discs obtained from postharvest tubers of treated plants. Thus the protective effect seemed to persist throughout the whole crop cycle. This treatment may offer the possibility of controlling both foliage and tuber blight and could have a major impact in reducing over-winter survival of P. infestans in tubers.

Enhancement of natural disease resistance in potatoes by chemicals.

Phosphites (Phi) are alkali metal salts of phosphorous acid, with the ability to protect plants against different pathogens. In this research, the effect of Phi applied to potato plants on severity of three important potato diseases in Argentina was assessed. Seed tubers and foliage of potato cvs Shepody and Kennebec were treated with Phi to assess effects on resistance against Phytophthora infestans, Fusarium solani and Rhizoctonia solani. Protection resulting from Phi treatment in seed tubers was high against P. infestans, intermediate against F. solani, and low against R. solani. In addition, seed tubers treated with calcium or potassium phosphites (CaPhi and KPhi, respectively) at 1% of commercial product emerged earlier than untreated ones. When Phi were foliarly applied two or four times at different doses, high levels of protection against P. infestans were achieved in both cultivars. Higher protection was observed in Kennebec when CaPhi was applied, while in Shepody this was true for KPhi. Expression of β-1,3-glucanases was induced at different times after treatment but no correlation between β-1,3-glucanases expression and foliar protection level was found. On the other hand, Phi positive protection effects did not produce negative effects in plant growth. Leaves from CaPhi-treated plants showed a darker green colour than leaves from control plants; also an increase in Rubisco protein and a delay in crop senescence was observed.

Phosphite compounds reduce disease severity in potato seed tubers and foliage

Although phosphite is widely used to protect plants from pathogenic oomycetes on a wide range of horticultural crops, the molecular mechanisms behind phosphite induced resistance are poorly understood. The aim of this work was to assess the effects of potassium phosphite (KPhi) on potato plant defense responses to infection with Phytophtora infestans (Pi). Pathogen development was severely restricted and there was also an important decrease in lesion size in infected KPhi-treated leaves. We demonstrated that KPhi primed hydrogen peroxide and superoxide anion production in potato leaves at 12 h post-inoculation with Pi. Moreover, the KPhi-treated leaves showed an increased and earlier callose deposition as compared with water-treated plants, beginning 48 h after inoculation. In contrast, callose deposition was not detected in water-treated leaves until 72 h after inoculation. In addition, we carried out RNA gel blot analysis of genes implicated in the responses mediated by salicylic (SA) and jasmonic acid (JA). To this end, we examined the temporal expression pattern of StNPR1 and StWRKY1, two transcription factors related to SA pathway, and StPR1 and StIPII, marker genes related to SA and JA pathways, respectively. The expression of StNPR1 and StWRKY1 was enhanced in response to KPhi treatment. In contrast, StIPII was down regulated in both KPhi- and water-treated leaves, until 48 h after infection with Pi, suggesting that the regulation of this gene could be independent of the KPhi treatment. Our results indicate that KPhi primes the plant for an earlier and more intense response to infection and that SA would mediate this response.

Gustavo Raúl Daleo

Papers

Accumulation of cytosolic glyceraldehyde-3-phosphate dehydrogenase RNA under biological stress conditions and elicitor treatments in potato.

Enhancement of natural disease resistance in potatoes by chemicals.

Phosphite compounds reduce disease severity in potato seed tubers and foliage

Potassium phosphite primes defense responses in potato against

Potassium phosphite primes defense responses in potato against Phytophthora infestans.