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Acetyl salicylic acid Aspirin and salicy
Plant Biology
University of the Punjab
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Plant Growth Regulation 30: 157–161, 2000. © 2000 Kluwer Academic Publishers. Printed in the Netherlands. 157
Acetyl salicylic acid (Aspirin) and salicylic acid induce multiple stress
tolerance in bean and tomato plants
Tissa Senaratna∗, Darren Touchell, Eric Bunn & Kingsley Dixon
Kings Park and Botanic Garden, West Perth, WA 6005, Australia ( ∗ author for correspondence; e-mail: tissa@kpbg.wa.gov)
Received 1 June 1999; accepted 24 August 1999
Key words: stress tolerance, acetyl salicylic acid, salicylic acid, heat, drought, cold
Abstract
The hypothesis that physiologically active concentrations of salicylic acid (SA) and its derivativescan confer stress tolerance in plants was evaluated using bean ( Phaseolus vulgaris L.) and tomato ( Lycopersicon esculentum L.). Plants grown from seeds imbibed in aqueous solutions (0–0 mM) of salicylic acid or acetyl salicylic acid (ASA) displayed enhanced tolerance to heat, chilling and drought stresses. Seedlings acquired similar stress tolerance when SA or ASA treatments were applied as soil drenches. The fact that seed imbibition with SA or ASA confers stress tolerance in plants is more consistent with a signaling role of these molecules, leading to the expression of tolerance rather than a direct effect. Induction of multiple stress tolerance in plants by exogenous application of SA and its derivatives may have a significant practical application in agriculture, horticultureand forestry.
Abbreviations: SA – salicylic acid; ASA – acetyl salicylic acid; SAR – systemic acquired resistance
1. Introduction
One of the most important factors that dictates the distribution of many plant species is their ability to withstand environmental stress including seasonal variations in temperature and available moisture. Plants generally respond to environmental stress by activating defence mechanisms and adjusting their cel- lular metabolism [8, 16]. Plants perceive the stress condition and signal to alter the metabolic flux for the activation/synthesis of defence mechanisms [16]. Many molecules, for example, calcium, jasmonic acid, ethylene and salicylic acid have been suggested as sig- nal transducers or messengers [11]. Salicylic acid (SA) has received much attention after the discovery of its ability to induce resistance (systemic acquired resist- ance or SAR) to pathogens [13, 19, 20, 25]. Exogen- ous application of SA induced pathogenesis-related gene expression and systemic acquired resistance [2]. Extensive studies have been undertaken to elucid- ate the molecular biology of SA induced SAR [2, 5,
17]. However, the physiological and biochemical basis for this phenomenon is not clear at present. Pathological disorders caused by microbial agents usually promote the development of hypersensitive reactions within the infected plant tissues. If the patho- gen is allowed to develop unchecked, necrotic lesions develop, resulting in cell and tissue death [1]. It has been demonstrated [10] that this sequence involves destructive attack by free radicals mediated through oxidative degradation of membrane lipids. Similarly there is considerable evidence to suggest that irreversible injury due to environmental stress is caused by increased free radical titre and consequent oxidation events which lead to degradation of bio- molecules such as membrane lipids and proteins [15, 22, 23]. The similarity of the injury mechanism between pathogenesis and stress leads us to hypothesise that salicylic acid which induces resistance to disease also confers tolerance to environmental stress. In this com- munication, evidence is provided that salicylic acid
Table 1. Survival (%) of SA or ASA – treated tomato and bean plants after heat, cold and drought stress
Heat Chilling Drought SI SD SI SD SI SD Conc(mM) Tomato Bean Tomato Bean Tomato Bean Tomato Bean Tomato Bean Tomato Bean SA0 0 00 00 00 00 00 0 0 000000000000 0 100 100 100 100 100 100 100 100 100 100 100 100 0 100 100 100 100 100 100 100 100 100 100 100 100 1 0 00 00 00 00 00 0
ASA0 0 00 00 00 00 00 0 0 000000000000 0 100 100 100 100 100 100 100 100 100 100 100 100 0 100 100 100 100 100 100 100 100 100 100 100 100 1 0 00 00 00 00 00 0 SI Seed Imbibed. SD Soil Drenched.
(SA) and acetyl salicylic acid (ASA) provide multiple stress tolerance in plants and that salicylic acid and its derivatives regulate the expression of stress toler- ance. This is the first report to suggest that SA or ASA imbibed into a seed impart tolerance to a variety of stresses in plants.
2. Materials and methods
To evaluate the hypothesis that SA and derivatives can confer stress tolerance, experimentation was conducted using bean ( Phaseolus vulgaris L. cv. brown beauty) and tomato ( Lycopersicon esculentum L. cv. romano) as test species. Plants were grown in 135 ml pots in a glasshouse maintained at ambient temperature and humidity (in Perth, Western Aus- tralia). Fourteen day-old plants were soil-drenched with 20 ml of distilled water or 0, 0, 0, 1. and 5 mM ASA or SA (dissolved in distilled water). Alternatively, seeds were imbibed in the solutions or in distilled water for 24 h and sown in pots. One week after soil-drenching or three weeks after the seed treatment, seedlings were subjected to heat, cold and drought stresses. For heat treatment, seed- lings were exposed to 54±0◦C for 3 h with an average light intensity of 40μMol m− 2 sec− 1 and then returned to room temperature. For chilling stress, plants were exposed to 0±0◦C in an incubator with an average light intensity of 35μMol m− 2 sec− 1 and 16/8 h light/dark photoperiod for two days. All pots were saturated with water 2 h prior to and after
the imposition of heat and chilling stress treatments. Drought stress was imposed by withholding water for 7 days, then on the 8th day all pots were watered until saturation. To evaluate the benefit of foliar spray applica- tions plants were sprayed with either 0 mM ASA or 0 mM SA, and subjected to stress treatments as above. The nozzle of the hand held sprayer was adjus- ted to deliver 0 ml of solution per spray and each plant received 1 ml of solution or distilled water for control. One week after spraying, plants were exposed to stress and scored for survival. All treatments were assessed after 48 h and sur- vival was determined by the ability of plants to regain turgidity and resume normal growth after the stress treatment. Damage was recorded as irreversible wilt- ing, desiccation of leaves and presence of necrotic leaf lesions. Randomized complete block design with 5 replicates was used in all experiments.
3. Results and discussion
Concentrations over 1 mM ASA or SA appeared to have adverse effects and therefore lower concentra- tions were used in further experiments. There were no observable differences in plant height, vigour or leaf number between non treated and SA or ASA treated plants (data not shown). Heat treatment at 54◦Cfor 3 h induced severe wilting in controls of tomato and bean plants and in plants treated with 0 mM SA or ASA. The plants, seed imbibed or drenched with
inhibit gibberellin biosynthesis in plants and hence have growth retarding effects. Salicylic acid and acetyl salicylic acid are not known to retard plant growth and no evidence of growth impairment of treated plants over untreated controls was observed in this study. The fact that seed imbibition of SA and ASA confers tolerance to plants suggests that these molecules trig- ger the expression of the potential to tolerate stress rather than having any direct effect as a protectant. Some reports suggest that SA induces an oxidative burst involving H 2 O 2 (hydrogen peroxide) accumu- lation which acts as the signal transducer for SAR [3]. Neuenschwander et al. [18] did not observe major changes in H 2 O 2 levels during the onset of SAR. The exact mechanism of action of SA in inducing SAR and its role in the transient increase of H 2 O 2 is still cause for debate [4]. Prevention of oxidative damage to cells during stress has been suggested as one of the mechanisms of stress tolerance [12, 24] and this level of protection is attributed to enhanced antioxid- ant activity [6, 21, 24]. Enhanced thermotolerance has been observed in potato microplants grown on media containing acetyl salicylic acid [14]. Increased ther- motolerance in mustard seedlings sprayed with SA has also been reported [7] and the tolerance was associ- ated with changes in antioxidants such as glutathione reductase, dehydroascorbate reductase and monode- hydroascorbate reductase [6]. In pea seedlings SA treatment decreased catalase and peroxidase levels with concomitant increase in glutathione reductase [26]. SA treatment also increased the level of reduced glutathione (GSH) with an increase in the ratio of reduced to oxidised glutathione (GSSG) indicating higher antioxidant potential [26]. There are sugges- tions that salicylic acid acts as an antioxidant [4]. However the fact that seed imbibition with SA or ASA provides stress tolerance in plants is more consistent with a signalling role for the expression of tolerance rather than a direct effect. The other mechanisms of tolerance may involve avoidance of lethal stress by altering cellular meta- bolism, synthesis of stabilising molecules to maintain integrity and function of cellular membranes during stress [27]. Stomatal resistance in SA treated mus- tard seedlings with enhanced thermotolerance was not different from controls [7]. However detailed studies on plant water relations after SA and ASA treat- ments are necessary to understand the events leading to stress tolerance. Although the physiological and biochemical basis for SA induced tolerance is not clearly understood, we believe that a cascade of events
is triggered to provide multiple stress tolerance in plants. Further research is warranted to elucidate the physiological and biochemical mechanisms by which SA induces tolerance to a variety of environmental stresses. In addition research is required to elucidate the reasons why plants treated with higher concentra- tions of SA or ASA (eg. 1 mM) were susceptible to stress injury. However, the phenomenon that common aspirin can be utilised to prevent crop losses during stress may have significant practical application.
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Acetyl salicylic acid Aspirin and salicy
Course: Plant Biology
University: University of the Punjab
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