Trends in Food Science & Technology 21 (2010) 502e509

Review

Impact of salicylic acid on post-harvest physiology of horticultural crops

Mohammadreza Asgharia,* and Morteza Soleimani Aghdamb
Department of Horticulture, Faculty of Agriculture, Urmia University, Nazloo street, Urmia, Iran (Tel.: D98 914 343 3494; fax: D98 441 297 2360; e-mails: m.asghari@urmia.ac.ir; mhamadreza@yahoo.com) b Iranian Young Researchers Club, Islamic Azad University, Ahar Branch, Ahar, Iran
Salicylic acid (SA), an endogenous plant growth regulator, has been found to generate a wide range of metabolic and physiological responses in plants thereby affecting their growth and development. SA as a natural and safe phenolic compound exhibits a high potential in controlling post-harvest losses of horticultural crops. In the present review, we have focused on various intrinsic biosynthetic pathways and effects of exogenous salicylic acid on post-harvest decay and disease resistance, oxidative stress, fruit ripening, ethylene biosynthesis and action, fruit rmness, respiration, antioxidant systems and nutritional quality have also been discussed.
a

been mainly on the roles of SA on biotic stresses, several studies also support major roles of salicylates in modulation of the plant response to several abiotic stresses, such as UV light, drought, salinity, chilling stress and heat shock (Ding & Wang, 2003; Ding, Wang, Gross, & Smith, 2001). Salicylates delay the ripening of fruits, probably through inhibition of ethylene biosynthesis or action, and maintain post-harvest quality (Srivastava & Dwivedi, 2000). Some of the results reported by several authors regarding the effects of SA on quality aspects of different harvested horticultural crops have been summarized in Table 1. Postharvest decay control in horticultural crops Plants continuously remain exposed to the challenging threats of a variety of pathogenic attacks. For many years synthetic fungicides were used to control post-harvest decay but, the public concerns about fungicide residues in fresh horticultural crops and the harmful effects of chemicals on human health and environment have recently caused scientists to search for new alternatives to chemical fungicides (Babalar, Asghari, Talaei, & Khosroshahi, 2007). Recent studies have shown that SA can be introduced as a potent alternative to chemicals (Table 1). Malamy, Carr, and Klessig (1990) showed that large amount of SA accumulates in the leaves of tobacco mosaic virus (TMV) resistant tobacco variety Nicotiana tabaccum cv. Xanthi upon inoculation with TMV. Further, SA or acetyl salicylic acid (ASA), a synthetic analogue of SA, when applied exogenously induced the expression of PR (pathogenesis related) genes and also conferred resistance against various pathogens (Malamy & Klessig, 1992). Exogenous application of SA at nontoxic concentrations to susceptible fruits and vegetables could enhance resistance to pathogens and control post-harvest decay (Asghari, Hajitagilo, & JaliliMarandi, 2009; Asghari, Hajitagilo, & Shirzad, 2007; Babalar et al., 2007). SA in a concentration dependent manner from 1 to 2 mmol L1 effectively reduced fungal decay in Selva strawberry fruit (Babalar et al., 2007). MeSA triggers disease resistance and mediates the expression of defense related genes in neighboring plants and in healthy tissue of infected plants (Shulaev, Silverman, & Raskin, 1997). In Hayward kiwifruit post-harvest decay was signicantly affected by MeSA vapor at the end of storage period. Decay incidence in fruit treated with 32 ml L1 was 6.3% whereas it was 34.2% in control fruits (Aghdam, Mosto, Motallebiazar, Ghasemneghad, & Fattahi

Introduction Salicylic acid (SA) and methyl salicylate (MeSA) are endogenous signal molecules, playing pivotal roles in regulating stress responses and plant developmental processes including heat production or thermogenesis, photosynthesis, stomatal conductance, transpiration, ion uptake and transport, disease resistance, seed germination, sex polarization, crop yield and glycolysis (Klessig & Malamy, 1994). Recently, SA has received a particular attention because it is a key signal molecule for expression of multiple modes of plant stress resistance. Although the focus has
* Corresponding author.
0924-2244/$ - see front matter 2010 Elsevier Ltd. All rights reserved. doi:10.1016/j.tifs.2010.07.009

M. Asghari, M.S. Aghdam / Trends in Food Science & Technology 21 (2010) 502e509 Table 1. Summary of some effects of SA on some harvested horticultural crops reported by different authors Author(s) Babalar et al. (2007) Sayyari, Babalar, Kalantari, Serrano, and Valero (2009) Zhang et al. (2003) Srivastava and Dwivedi (2000) Wang, Chen, Kong, Li, and Archbold (2006) Yao and Tian (2005) Fung et al. (2004) Ding and Wang (2003) Ding et al. (2001) Hung, Liu, Lu, and Xia (2007) Mo et al. (2008) Aghdam et al., (2009) Fung et al. (2006) Xu and Tian (2008) Jing-Hua, Yuan, Yan-Man, Xiao-Hua, and Zhang (2008) Yu and Zheng (2006) Shaee, Taghavi, and Babalar (2010) Cao, Zeng, and Jiang (2006) Cai et al. (2006) Peng and Jiang (2006) Reported results Marketability retention, decrease in ethylene production & fungal decay Inhibition of PAL activity, retention of vitamin C content reduction of CI & EL Inhibition of ACS, ACO & LOX activity, suppression of ethylene & superoxide free radical production, increase in total SA content, Decrease in fruit softening, pulp/peel ratio,reducing sugar content, invertase activity & respiration rate, inhibition of cellulase, PG, xylanase, CAT & POX activity Increase in rmness, APX, GR activity, AsA/DHAsA & GSH/GSSG ratios, decrease in CI, DI induction of HSP101 & HSP73 genes Increase in b-1, 3-glucanase, PAL & POD activity, induction of disease resistance, direct antifungal activity Increase in transcript levels of AOX, CI protection Development of red color, regulation of ACS genes expression Accumulation of HSPs, CI reduction Acceleration of H2O2 accumulation, increase in SOD, GR, APX & DHAR activity & ASA/ DHASA & GSH/GSSG ratios, decrease in lipid peroxidation & MDA Decreased ethylene, LOX activity, MDA content, DI, softening & respiration rates, TSS & activated the SOD, POD, CAT&APX enzymes Inhibition of ethylene production, ripening & decay control Expression of AOX, CI resistance Decay control, increase in CAT, GPX, b-1, 3-glucanase& chitinase Increase in GPX, APX, CAT, SOD & GR activity, induction of resistance to CI Enhanced the biological efcacy of antagonist C. laurentii Decay, Ripening& weight loss reduction Inhibition of PAL, CAT & POD decrease in superoxide free radical production & CI Induction of resistance to diseases, increase in POD, PAL, GR, Chitinase & b-1, 3-glucanase, decrease in CAT & APX Delayed discoloration, maintained edible quality, activity of PPO, POD & PAL

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Commodity Strawberry Pomegranate Kiwifruit Banana Peach Sweet cherry Sweet pepper Tomato Tomato Orange Apple Kiwifruit Tomato Sweet cherry Watermelon Apple Strawberry Loquat Pear Fresh-cut Chestnut

DI decay index, GR glutathione reductase, GSSG oxidized glutathione, GSH reduced glutathione, DHAR dehydroascorbate reductase, AsA ascorbate, DHAsA dehydroascorbate, MDA Malondialdehyde.

Moghaddam, 2009). Dipping of pear fruit in 1 mmol L1 SA solution effectively controlled fruit decay during 5 months of cold storage (Asghari et al., 2007). Treatment of strawberry plants at vegetative stage and fruit development stage followed by post-harvest treatment of fruits with 1 and 2 mmol L1 effectively controlled the total decay and increased fruit shelf life (Babalar et al., 2007). Postharvest treatment of table grapes with SA before coating with chitosan signicantly enhanced the efcacy of coating and decreased fruit total decay (Asghari et al., 2009). Some researches indicate that SA also exhibits direct antifungal effects against pathogens. Lu and Chen (2005) have demonstrated the inhibitory action of SA on botrytis rot in lily leaves. Foliar application of Asilbenzolar-S-Methyl (a synthetic analogue of SA) has led to protection of post-harvest Rock melons and Hami melons from diseases (Huang, Deverall, Tang, Wang, & Wu, 2000). 2 mmol L1 SA showed direct fungal toxicity on Monilinia fructicola and signicantly inhibited the mycelial growth and spore germination of the pathogen in vitro (Yao & Tian, 2005). SA also effectively enhances the biocontrol efcacy of antagonist yeasts. According to the results of Qin, Tian,

Xu, and Wan (2003), 0.5 mmol L1 SA signicantly reduced the incidence of blue mould (P. expansum) and alternaria rot (A. alternata) in sweet cherry without any surface injury. Adding SA signicantly improved the activity of R. glutinis against both pathogens. Plants protect themselves against the pathogen attacks by activating some kinds of defense mechanisms such as local acquired resistance (LAR) and systemic acquired resistance (SAR) (Vlot, Dempsey, & Klessig, 2009). As seen in Fig. 1, salicylates are a major component in the signal transduction pathways of plants playing an important role in disease resistance (Park, Kaimoyo, Kumar, Mosher, & Klessig, 2007). Once plant defense responses are activated at the site of infection (LAR), a systemic defense response is often triggered in distal plant parts to protect these undamaged tissues against subsequent invasion by the pathogen. This long-lasting and broad-spectrum induced disease resistance is referred to as SAR and is characterized by the coordinate activation of a specic set of PR-genes, many of which encode for proteins with antimicrobial activity (Durrant & Dong, 2004; Van Loon, Rep, & Pieterse, 2006). The onset of SAR is associated with increased levels

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rate of superoxide radical (O2) generation in SA-treated fruits were higher than that in controls after 8 days of treatment. According to the ndings of Tian et al. (2007) the balance between superoxide dismutase (SOD), POD and catalase (CAT) activities in cells is crucial for determining the steady-state level of O2 and H2O2. SA interaction with the above mentioned enzymes leads to high levels of H2O2 accumulating in cells, which induces fruit resistance against pathogens via activating protective enzymes and PR-proteins (Klessig & Malamy, 1994; Malamy & Klessig, 1992;). It has been demonstrated that, SA inhibits the H2O2-scavenging activity of cytosolic ascorbate peroxidase (APX), and H2O2 levels concomitantly rise upon SA treatment of tobacco leaves (Fig. 1). Thus, SA may also facilitate H2O2 accumulation during the oxidative burst (OB) induced by infection with the virulent pathogens. The increased ROS associated with the OB may contribute to resistance via several mechanisms, including directly killing the invading pathogen and/or activating cell wall crosslinking and lignication, thereby strengthening the cell wall and helping conne the pathogen to the infection site (Dempsey, Shah, & Klessig, 1999) SA prevents post-harvest oxidative stress and alleviates chilling injury during cold storage Plant defense system against oxidative stress consists of two lines; The rst line of defense is termed ROS avoidance genes includes alternative oxidase (AOX) and the second is termed as ROS scavenging genes includes SOD, CAT, the ascorbate/glutathione cycle, the glutathione peroxidase system and thioredoxin system (Fig. 2) (Buchanan, Gruissem, & Jones, 2000, p 1343). SA has been shown to induce expression of AOX and increase the antioxidant capacity of the cells (Table 1, Fig. 2). For example, SA stimulates the synthesis of antioxidant enzymes and induces the activity of PPO, PAL and b-1, 3-glucanase in sweet cherries (Fig. 3) (Qin et al., 2003). Yao and Tian (2005) reported that pre-harvest treatment of sweet cherries with SA has induced b-1, 3-glucanase, PAL and POD activities during the short time storage period and the activity of these enzymes in SA-treated cherries stored at 25  C was higher than in fruits stored at 0  C. SA, in a concentration dependent manner from 0 to 2 mmol L1, enhanced the strawberry fruit total antioxidant capacity (TAC). 2 mmol L1 was the most effective concentration while SA at 4 mmol L1 caused a slight increase in fruit TAC. Consequent application of SA at three stages of vegetative growth, fruit development and post-harvest stage was the most effective strategy in improving fruit TAC (Asghari, & Babalar, 2009). Postharvest treatment of sweet cherry fruits with SA signicantly inhibited CAT activity, but stimulated the activity of SOD and POD. After inoculation with P. expansum, CAT activity decreased and SOD activity increased in SA-treated fruits. SA treatment also changed the expression of POD isozymes, indicating that SA directly or indirectly activates antioxidant enzymes (Tian et al., 2007).

Fig. 1. SA decreases the expression of ascorbate peroxidase (APX) & catalase (CAT) genes leading to substantial increase in H2O2 which is crucial for the activation of local acquired resistance (LAR) & systemic acquired resistance (SAR).

of SA, locally at the site of infection and often also systemically in distant tissues (Klessig & Malamy, 1994; Malamy & Klessig, 1992; Tsuda, Sato, Glazebrook, Cohen, & Katagiri, 2008). Park et al., (2007) demonstrated that MeSA acts as a crucial long distance SAR signal in tobacco. SA can induce the accumulation of hydrogen peroxide (H2O2) levels in plant tissues which acts as a signal activating the SAR (Fig. 1) (Klessig & Malamy, 1994; Tian, Qin, Li, Wang, & Meng, 2007). Mutant and transgenic plants that are impaired in SA signaling are incapable of developing SAR and dont show PR gene activation upon pathogen infection, which indicates that SA is a necessary intermediate in the SAR signaling pathway (Durrant & Dong, 2004; Pieterse, Leon-Reyes, Van der Ent, & Van Wees, 2009). The SA induced defense responses are probably involved in the expression of a range of defense genes, especially those encoding PR-proteins such as chitinase, b-1, 3-glucanase and peroxidase (POD) (Meena, Marimuthu, & Velazhahan, 2001). According to Zeng, Cao, and Jiang (2006), in a study on mangoes after 4 days of treatment the activity of b-1, 3-glucanase in SA-treated fruits were over higher than in controls. They found that the level of hydrogen peroxide (H2O2) and the

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Fig. 2. Antioxidant systems in plants.

Chilling injury (CI) is a type of damage caused by low temperature as a result of oxidative burst. Although there are many methods to reduce CI in various horticultural crops, SA and MeSA treatments are inexpensive, easy to set up and applicable to various horticultural crops (Ding et al., 2001). Increase in AOX transcript levels using SA and MeSA before cold treatment reduced incidence of CI in freshly green bell peppers (Fung, Wang, Smith, Gross, & Tian, 2004). In response to abiotic stresses, all living organisms synthesis new proteins, for example, plants respond to high temperatures with the synthesis of a group of proteins known as heat shock proteins (HSPs) (Fig. 3). Often the accumulation of HSPs not only confers protection against the stress that causes their biosynthesis but also against any other subsequent stress situation (Tian et al., 2007). Treatments with SA and MeSA prior to low-temperature storage induce HSPs biosynthesis and, at the same time, CI tolerance in tomatoes and peaches (Ding et al., 2001). Accumulation of the HSPs in chilling-sensitive horticultural products with SA and MeSA treatments would

allow their storage at low temperatures without CI development. On the other hand lipid peroxidation is one of the adverse effects of CI on plant cells leading to malondialdehyde (MDA) accumulation. It has been reported that MDA accumulation is prevented after SA treatment (Table 1 & Fig. 3) SA delays fruit ripening Impact of SA on fruit softening Fruit ripening and senescence are accompanied by changes in several quality aspects such as softening, decrease in total acidity and increase in sugar contents, color development, aroma production and etc. (Wills, McGlasson, Graham, & Joyce, 1998). Softening of fruits is a main and critical quality change. MeSA, in a concentration dependent manner from 0 to 32 ml L1, maintained rmness of kiwifruit during storage (Aghdam et al., 2009). Srivastava and Dwivedi (2000) reported that in bananas treated with SA fruit softening markedly decreased. Zhang, Chen, Zhang, and Ferguson (2003)

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Fig. 3. Induction of chilling resistance in plants by SA.

reported a positive correlation between fruit free SA content and rmness in kiwifruit during ripening. Retention of fruit rmness as the result of SA treatment has been reported in several crops (Table 1). It has been demonstrated that SA decreases ethylene production and inhibits cell wall and membrane degrading enzymes such as polygalacturonase (PG), lipoxygenase (LOX), cellulase and pectinemethylesterase (PME) leading to decreasing the fruit softening rate (Srivastava & Dwivedi, 2000; Zhang et al., 2003). Effect on TSS and sugars TSS and soluble sugars may increase during fruit ripening due to the action of sucrose-phosphate synthase (SPS), a key enzyme in sucrose biosynthesis (Hubbard, Pharr, & Huber, 1991). This enzyme is activated by ethylene and the ripening process itself during storage (Langenkamper, McHale, Gardner, & MacRae, 1998). Recently, an increase in sucrose-phosphate synthase and invertase activities and a decrease in sucrose synthase activity have been reported during ripening of some fruits (Cordenunsi & Lajolo, 1995). Treatment of kiwifruits with MeSA of 32 ml L1 maintained a lower TSS content than the control fruits at the end of cold storage (Aghdam et al., 2009). The authors proposed that MeSA reduced ethylene production may results to decreased SPS enzyme activity leading to decrease in sucrose synthesis. The ripening of banana fruit is accompanied by increase in pulp to peel ratio. Rise in pulp to peel ratio during fruit ripening may be due to change in sugar concentration in the two tissues. A rapid increase in sugar contents in the pulp

than those in the peel leads to a change in osmotic pressure, as a result of which water is withdrawn from the peel and hence pulp to peel ratio increases accordingly. According to the ndings of Srivastava and Dwivedi (2000), SA treatment, in a concentration dependent manner, reduces this increase in pulp to peel ratio, leading to a delay in banana fruit ripening. The result showed that the invertase activity is concomitant with decrease in nonreducing sugar content. The level of reducing sugars is increased and nonreducing sugars is decreased during ripening and senescence. This accumulation of reducing sugars may be due to increased breakdown of starch during ripening as reported by Beaudry, Ray, Clanton, and Stanley (1989). SA treatment resulted in decreased levels of invertase and reducing sugar contents while an opposite effect on nonreducing sugar content, suggesting that SA delays banana fruit ripening. On the other hand cell walls contain large amounts of polysaccharides, mainly pectins and cellulose, and are digested due to the activity of the cell wall degrading enzymes leading to a signicant increase in TSS content. Then any factor preventing these enzymes will prevent from a dramatic increase in TSS content. As seen in Table 1 and Fig. 4, SA effectively protects cell walls by decreasing the expression of degrading enzymes and as a consequence prevents from dramatic increase in TSS content of the cells. Inhibition of ethylene biosynthesis Ethylene plays a key role in fruit ripening and senescence. This hormone triggers the induction of cell wall hydrolyzing enzymes leading to increase in respiration rate, fruit softening and senescence (Wills et al., 1998). SA

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Fig. 4. Schematic overview of mechanisms by which SA delays fruit ripening and extends storage life.

effectively decreases ethylene production in several horticultural crops (Table 1). It has been shown that MeSA treatment signicantly decreases ethylene production in kiwifruits (Aghdam et al., 2009). Both SA and ASA have been shown to inhibit ethylene production in cultured pear cells, mung bean hypocotyls, apple and pear fruit tissue discs, carrot cell suspension cultures and some fruits (Babalar et al., 2007; Romani, Hess, & Leslie, 1989). Srivastava and Dwivedi (2000) reported that SA has delayed the ripening of banana fruit, probably through inhibition of ethylene biosynthesis or action. SA decreases ethylene production by decreasing ACS and ACO production and activity. Zhang et al., (2003) reported that postharvest treatment of kiwifruit with ASA resulted in a lower ACO and ACS activity and decreased ethylene production during the early stages of fruit ripening. Effect on LOX activity There is evidence for a positive correlation between LOX activity, free radicals production and ethylene biosynthesis in fruit tissue (Marcelle, 1991). SA inhibited woundinduced transcription and also activity of ACS in tomato and decreased LOX activity in disks of kiwifruit leading to the consequent reduction in the production of free radicals and ethylene biosynthesis (Zhang et al., 2003). Ding and Wang (2003) showed that ripening process in mature green tomatoes was enhanced by 0.1 mmol L1 MeSA and by 0.01 mmol L1 during breaker stage. But in fruit at turning stage even 0.01 mmol L1 SA inhibited the ripening process. 0.5 mmol L1 SA prevented red color

development and increased ethylene production and respiration rate in all maturity stages. Effect on cell respiration SA has been reported to effectively reduce the respiration rate in several fruits (Table 1). SA as a signal triggers the induction of cyanide resistance respiration in plant cells by affecting the AOX enzyme activity (Raskin, Turner, & Melander, 1989). In horticultural crops, SA affects AOX activity leading to decrease in the harmful effects of different post-harvest oxidative stresses such as chilling injury, prevents fermentation, and maintains low respiration rates and decreases fruit ripening and senescence rates. Respiration of harvested crops is highly dependent to ethylene production and activity and any factor increasing the production and activity of ethylene leads to increases in respiration and consequently increases the senescence rate. Effect of SA in decreasing the respiration rate is mainly due to its negative effects on ACC, ACO, PG, PME, cellulase and antioxidant enzymes leading to decrease in ethylene production and action (Fig. 4). Conclusion SA, as a natural and safe phenolic compound, exhibits a high potential in controlling post-harvest losses of horticultural crops. Decrease in ethylene production and action, induction of disease resistance, prevention of oxidative stresses, induction of crop tolerance to chilling injury, decrease in respiration rate, decrease in ripening and senescence rate, prevention of cell wall degrading enzymes and

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M. Asghari, M.S. Aghdam / Trends in Food Science & Technology 21 (2010) 502e509 Ding, C. K., Wang, C. Y., Gross, K. C., & Smith, D. L. (2001). Reduction of chilling injury and transcript accumulation of heat shock protein genes in tomatoes by methyl jasmonate and methyl salicylate. Plant Science, 161, 1153e1159. Durrant, W. E., & Dong, X. (2004). Systemic acquired resistance. Annual Review of Phytopathology, 42, 185e209. Fung, R. W. M., Wang, C. Y., Smith, D. L., Gross, K. C., Tao, Y., & Tian, M. (2006). Characterization of alternative oxidase (AOX) gene expression in response to methyl salicylate and methyl jasmonate pre-treatment and low temperature in tomatoes. Journal of Plant Physiology, 163, 1049e1060. Fung, R. W. M., Wang, C. Y., Smith, D. L., Gross, K. C., & Tian, M. (2004). MeSA and MeJA increase steadyestate transcript levels of alternative oxidase and resistance against chilling injury in sweet peppers. Plant Science, 166, 711e719. Huang, Y., Deverall, B. J., Tang, W. H., Wang, W., & Wu, F. W. (2000). Foliar application of asilbenzolar-S-methyl and protection of postharvest rock melons and Hami melons from disease. European Journal of Plant Pathology, 106, 651e656. Hubbard, N. L., Pharr, D. M., & Huber, S. C. (1991). Sucrose phosphate synthase and other sucrose metabolizing enzymes in fruits of various species. Plant Physiology, 82, 191e196. Hung, R. H., Liu, J. H., Lu, Y. M., & Xia, R. X. (2007). Effect of salicylic acid on the antioxidant system in the pulp of Cara cara navel orange (Citrus sinensis L. Osbeck) at different storage temperatures. Postharvest Biology and Technology, 47, 168e175. Jing-Hua, Y., Yuan, G., Yan-Man, L., Xiao-Hua, Q., & Zhang, M. F. (2008). Salicylic acid-induced enhancement of cold tolerance through activation of antioxidative capacity in watermelon. Scientia Horticulture, 118, 200e205. Klessig, D. F., & Malamy, J. (1994). The salicylic acid signal in plants. Plant Molecular Biology, 26, 1439e1458. Langenkamper, G., McHale, R., Gardner, R. C., & MacRae, E. (1998). Sucrose phosphate synthase steady-state mRNA increases in ripening kiwifruit. Plant Molecular Biology, 36, 857e869. Lu, Y. Y., & Chen, C. Y. (2005). Molecular analysis of lily leaves in response to salicylic acid effective towards protection against Botrytis elliptica. Plant Science, 169, 1e9. Malamy, J., Carr, J. P., & Klessig, D. F. (1990). Salicylic acid: a likely endogenous signal in the resistance response of tobacco to viral infection. Science, 250, 1002e1004. Malamy, J., & Klessig, D. F. (1992). Salicylic acid and plant disease resistance. Plant Journal, 2, 643e654. Marcelle, R. D. (1991). Relationship between mineral content, lipoxygenase activity, levels of 1-aminocyclopropane-l-carboxylic acid and ethylene emission in apple fruit esh disks during storage. Postharvest Biology and Technology, 1, 101e109. Meena, B., Marimuthu, T., & Velazhahan, R. (2001). Salicylic acid induces systemic resistance in groundnut against late leaf spot caused by Cercosporidium personatum. Journal of Mycology and Plant Pathology, 31, 139e145. Mo, Y., Gong, D., Liang, G., Han, R., Xie, J., & Li, W. (2008). Enhanced preservation effects of sugar apple fruits by salicylic acid treatment during postharvest storage. Journal of the Science of Food and Agriculture, 88, 2693e2699. Park, S. W., Kaimoyo, E., Kumar, D., Mosher, S., & Klessig, D. F. (2007). Methyl salicylate is a critical mobile signal for plant systemic acquired resistance. Science, 318, 113e116. Peng, L., & Jiang, Y. (2006). Exogenous salicylic acid inhibits browning of fresh-cut Chinese water chestnut. Food Chemistry, 94, 535e540. Pieterse, C. M. J., Leon-Reyes, A., Van der Ent, S., & Van Wees, S. C. M. (2009). Networking by small-molecule hormones in plant immunity. Nature Chemical Biology, 5, 308e316. Qin, Q. Z., Tian, S. P., Xu, Y., & Wan, Y. K. (2003). Enhancement of biocontrol efcacy of antagonistic yeasts by salicylic acid in sweet cherry fruit. Physiological and Molecular Plant Pathology, 62, 147e154.

maintaining crop rmness are of main results obtained following SA treatment. The application of SA and especially pre-harvest, for inducing the defense resistance systems against post-harvest diseases may be a useful and promising measure for controlling post-harvest decay on a commercial scale. Since it effectively enhances the effects of other postharvest treatments, such as heat treatments and biocontrol agents, use of SA in combination with other post-harvest treatments may give better results in controlling post-harvest losses. SA can be used as an appropriate alternative to chemicals in post-harvest technology of horticultural crops to assure food safety. Since SA, like any other post-harvest treatment, may have different effects on different crops at different circumstances, it is necessary to determine the best and safe concentration for each crop cultivar.

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