تنش سرما و اثر آن بر سنتز هورمون اسید آبسیزیک از طریق بررسی بیان ژن و تغییرات آنتیاکسیدانهای غیرآنزیمی گوجه فرنگی (Solanum lycopersicum 'Red Cloud)
الموضوعات :نیلوفر صمدی 1 , سکینه سعیدی سار 2 , حسین عباسپور 3 , ناهید مسعودیان 4
1 - گروه زیست شناسی، واحد دامغان، دانشگاه آزاد اسلامی، دامغان، ایران
2 - گروه کشاورزی، دانشکده فنی و حرفه ای دکتر شریعتی، دانشگاه فنی و حرفه ای، تهران، ایران
3 - گروه زیست شناسی، واحد دامغان، دانشگاه آزاد اسلامی، دامغان، ایران
4 - گروه زیست شناسی، واحد دامغان، دانشگاه آزاد اسلامی، دامغان، ایران
الکلمات المفتاحية: آبسیزیک اسید, آنتیاکسیدانهای غیرآنزیمی, ریل تایم پی سی آر, SlNCED1, Solanom lycopersicum L,
ملخص المقالة :
گوجهفرنگی یکی از محصولات عمده کشاورزی در ردهی محصولات زراعی است که در بسیاری از کشورهای جهان و همچنین ایران از جنبههای مختلف دارای اهمیت میباشد. در این تحقیق تحت تنش سرما به بیان ژن SlNCED1در مسیر بیوسنتز هورمون آبسیزیک اسید به روش Real-Time qRT-PCR و اندازهگیری برخی خواص فیزیولوژیکی پرداخته شده است. بدینمنظور پس از میوهدهی، بوتهها بهمدت 12 و 24 ساعت در دمای 2 و 4 درجه سانتیگراد قرار گرفتند. پس از این مدت میوهها جهت سنجش آنتی اکسیدانهای غیرآنزیمی به روش DPPD و فنل کل به روش فولین سیکالتو و بیان ژن جدا شدند. نتایج حاکی از آن بود که بیشترین میزان بیان ژن SlNCED1مربوط به دمای 4 درجه بهترتیب در ساعات 24 و12 بود. همچنین نتایج آنالیز مقایسه میانگین نشان داد که فعالیت معنی دار آنتی اکسیدان وفنل کل در دمای 2درجه و 12 ساعت اتفاق افتاد. بهطورکلی میتوان گفت که از آنجایی که گوجه فرنگی رد کلود متعلق به نواحی گرمسیری است لذا در دماهای پایینتر از 12 درجه سانتیگراد دچار تنش سرمایی شده و متعاقب آن تغییرات بیان ژنها و آنتی اکسیدانها را جهت حفاظت از خود در برابر تنش ایجاد میکند.
Asadi, S., Moghaddam, M., Ghasemi pirbalouti, A. and Fotovat, A. (2018). Evaluation of physiological characteristics and antioxidant activity of sweet basil (Ocimum basilicum cv. Keshkeni luvelou) under different levels of methyl jasmonate and lead toxicity. Journal of Iranian Plant Ecophysiological Research. 13(51): 1-16.
Alscher, R.G. and Cumming, J.R. (1990). Stress responses in plants: adaptation and acclimation Mechanisms. John Wiley and Sons. INC Publication.
Banerjee, S., Yuan, X., Germida, J.J. and Vujanovic V. (2014). Gene expression patterns in wheat coleorhiza under cold-and biological stratification. Microbiological research. 169(7-8): 616-622.
Bose, J., Rodrigo-Moreno A. and Shabala, S. (2014). ROS homeostasis in halophytes in the context of salinity tolerance. Journal of Experimental Botany. 65: 1241–1257.
Brand-Williams, W., Cuvelier, M.E. andBerset, C. (1995). Use of free radical method to evaluate antioxidant activity. LWT- Food Science and Technology. 28(1):25-30
Bunn, R., Lekberg, Y. and Zabinski, C. (2009). Arbuscular mycorrhizal fungi ameliorate temperature stress in thermophilic plants. Ecology. 90:1378–1388.
Davies, P.J. and Mansfield, TA. (1983). The role of abscisic acid in drought avoidance, In: Abscisic acid (ed. by F.T. Adicott). Praeger, New York, 237-268.
Elhamirad, A.H. and Zamanipoor, M.H. (2012). Thermal stability of some flavonoids and phenolic acids in sheep tallow olein. European Journal of Lipid Science and Technology. 114(5): 602-606.
Ghorbanli, M., Rostami Abousaeidi, M. and Bakhshi Khaniki, Gh.R. (2014). Effect of naphthalene acetic acid (NAA) and abscisic acid (ABA) on quality and quantity of Strawberry Selva cultivar (Fragaria ananassa cv. Selva) and Paros cultivar (Fragaria ananassa cv. Paros). Journal of Iranian Plant Ecophysiological Research. 9: 30-39.
Großkinsky, D.K., Naseem, M., Abdelmohsen, U.R., Plickert, N., Engelke. T., Griebel, T., Zeier. J., Novák, O., Strnad, M., Pfeifhofer, H., Graaff, E.V.D. Simon, U. and Roitsch, T. (2011). Cytokinins mediate resistance against Pseudomonas syringaein tobacco through increased antimicrobial phytoalexin synthesis independent of salicylic acid signaling. Plant Physiol, 157:815–830
Hiwasa, K., Kinugasa, Y., Amano, S., Hashimoto, A., Nakano, R., Inaba, A. and Kubo, Y. (2003). Ethylene is required for both the initiation and progression of softening in pear Pyrus communis L. fruit. Journal of Experimental Botany. 54: 771–779.
Hosseinzad, B.E., Chaparzadeh N. and Dilmaghani K. (2014). Effect of salicylic acid on growth parameters, osmolytes and osmotic potential in radish (Raphanus sativus L.) under salt stress. Journal of Plant Research. 27(1): 32-40. (In Persian)
Huber, D.J. (1983). Polyuronide degradation and hemicellulose modification on ripening tomato fruit. Journal of the American Society for Horticultural Science. 108: 405–409.
Jahan tigh haghighi, Z., Fahmideh, L., Fazeli nasab, B. (2018). Evaluation and comparison of Leaf antioxidant properties and morphological traits of tomato varieties (Lycopersicon esculentum L). 13(50): 63-76.
Jajic, I., Sarna, T. and Strzalka, K. (2015). Senescence, Stress, and Reactive Oxygen Species. Plants. 4(3): 393-411.
Jamshidi, M., Ahmadi, H.R., Rezazadeh, Sh., Fathi, F. and Mazanderani, M. (2010). Study on phenolicd and anioxidant activity of some selected plant of Mazandaran province. Medic Plan. 934:177-183. (In Persian).
Ji, K., Kai, W., Zhao, B., Sun, Y. and Yuan, B. (2014). SlNCED1and SlCYP707A2: key genes involved in ABA metabolism during tomato fruit ripening. Journal of Experimental Botany. 6518: 5243-5255.
Kilian, J., Whitehead, D., Horak, J., Wanke, D., Weinl, S., Batistic, O.D., Angelo, C., Bornberg-Bauer, E., Kudla, J. and Harter, K. (2007). The AtGen Express global stress expression data set: protocols, evolution and model data analysis of UV-B light, drought and cold stress responses. The Plant Journal. 50:347–363.
Kok, E., Islek, C. and Sulun Ustun A. (2010). Effect of Cold on Protein, Proline, Phenolic Compounds and Chlorophyll Content of Two Pepper Capsicum annuum L. Varieties. G.U. Journal of Science. 231: 1-6.
Leon, P. and Sheen, J. (2003). Sugar and hormone connections. Trends Plant Science. 8: 110-116.
Liang, L.H., Mei, X., Lin, F., Xia, J., Liu, S.J. and Wang, J.H. (2009). Effect of low temperature stress on tissue structure and physiological index of cashew young leaves. Journal of Ecology and Environmental Sciences. 18: 317-320.
Mohammadi, B., Zia Jahromi, N., Sadeghi, H. and Mirzaei, A. (2016). Compare the amount of phenols, flavonoids and antioxidant activity of five varieties of Iranian olive leaf hydroalcoholic extract. Bringing Knowledge, 20 (10): 888-898. (In Persian).
Munder, M. (2009). Arginase: an emerging key player in the mammalian immune system: review. British Journal of Pharmacology. 158: 638–651.
Nakamura, S., Chono, M., Abe, F. and Miura, H. (2010). Mapping a diploid wheat abscisic acid 8′-hydroxylase homologue in the seed dormancy QTL region on chromosome 5Am. Euphytica. 171(1):111.
Oliver, S.N., Dennis, E.S. and Dolferus, R. (2007). ABA regulates apoplastic sugar transport and is a potential signal for cold-induced pollen sterility in rice. Plant and Cell physiology. 48(9): 1319-1330.
Pakkish, Z., Rahemi, M. and Baghizadeh, A. (2009). Seasonal changes of peroxidase, polyphenol oxidase enzyme activity and phenol content during and after rest in pistachio Pistacia vera L. Flower Buds. World Applied Sciences. 69: 1193-1199.
Qin, X.Q. and Zeevaart J.A. (1999). The 9-cis-epoxycarotenoid cleavage reaction is the key regulatory step of abscisic acid biosynthesis in water-stressed bean. Proceedings of the National Academy of sciences. 96(26): 15354-15361.
Qin, X.Q. and Zeevaart, J.A.D. (2002). Overexpression of a 9-cis-epoxycarotenoid dioxygenase gene in Nicotiana plumbaginifolia increases abscisic acid and phasic acid levels and enhances drought tolerance. Plant Physiology. 128: 544–551.
Raghavendra, A.S., Gonugunta, V.K., Christmann, A. and Grill, E. (2010). ABA perception and signaling. Trends Plant Science. 15:395–401.
Rivero, R.M., Ruiz, J.M., Garcıa, P.C., Lopez-Lefebre, L.R., Sanchez, E. and Romero, L. (2001). Resistance to cold and heat stress: accumulation of phenolic compounds in tomato and watermelon plants. Plant science. 160: 315-321.
Salmanian, S., Sadeghi Mahoonak, A. Alami, M. and Ghorbani, M. (2014). Evaluation of Total Phenolic, Flavonoid, Anthocyanin Compounds, Antibacterial and Antioxidant Activity of Hawthorn (Crataegus Elbursensis) Fruit Acetonic Extract. Journal of Rafsanjan University of Medical Sciences. 13 (1): 53-66. (In Persian)
Sangwan, V. and Dhindsa, R.S. (2002). In vivo and in vitro activation of temperature-responsive plant map kinases. FEBS Letters. 531:561–564.
Sayyari, M., Babalar, M., Kalantari, S., MartinezRomero, D., Guillen, F., Serrano, M. and Valero, D. (2011). Vapour treatments with methyl salicylate or methyl jasmonate alleviated chilling injury and enhanced antioxidant potential during postharvest storage of pomegranates. Food Chemistry. 124: 964–970.
Schlegl, Z. (1996). Photo assimilate distribution in plants and crops, source-sink relationships. Environmental Control in Biology. 323-330.
Schwartz, S.H., Qin, X.Q. and Zeevaart, J.A.D. (2003). Elucidation of the indirect pathway of abscisic acid biosynthesis by mutants, genes, and enzymes. Plant Physiology. 131:1591–1601.
Seppanen, M.M. (2000). Characterize of freezing tolerance in Solanum commersoniidun. with special reference of the relationship between and oxidative stress. University of Helsinki department of production section of crop husbandry. 56:4-44.
Sun, L., Wang, Y.P., Chen, P., Ren, J., Ji, K., Li, Q., Li, P., Dai, S.J. and Leng, P. (2011). Transcriptional regulation of SlPYL, SlPP2C, and SlSnRK2 gene families encoding ABA signal core components during tomato fruit development and drought stress. Journal of Experimental Botany. 62:5659–5669.
Taffazoli, A.A., and Kalabil, R. (2002). Increasing sugars and cold resistance in Kiwi fruit by ABA hormone. Iran Agricultural Research. 11. (In Persian)
Taylor, I.B., Sonneveld, T., Bugg, T.D.H. and Thompson, A.J. (2005). Regulation and manipulation of the biosynthesis of abscisic acid, including the supply of xanthophyll precursors. Journal of Plant Growth Regulation. 24:253–273.
Van kiet, H., Nose, A. and Shao, H. (2016). Effect of Cold Stress on Root Growth, Accumulation of Soluble Proteins and Free Amino Acids of Sheath Blight-Resistant Rice Genotype 32R. Tropical Agriculture and Development. 60(3): 191-194.
Wang, G.D., Liu, Q., Shang, X.T., Chen, C., Xu, N., Guan, J. and Meng, Q.T. (2017). Overexpression of transcription factor SlNAC35 enhances the chilling tolerance of transgenic tomato. Biologia Plantarum. 62(3):479–488.
Weidner, S., Kordala, E., Brosowska-Arendt, W., Karamac, M., Kosinska, A. and Amarowicz, R. (2009). Phenolic compounds and properties of antioxidants in grapevine roots Vitis vinifera L. under low temperature stress followed by recovery. Acta Societatis Botanicorum Poloniae. 78: 279-286.
Went, F.W. (1953). The effect of temperature on plant growth. Annual Review of Plant Physiology. 4:347–362.
Xiong, L., Wangm R.G., Mao, G. and Koczan, J.M. (2006). Identification of drought tolerance determinants by genetic analysis of root response to drought stress and abscisic acid. Plant Physiology. 142: 1065–1074.
Zhang, M., Yuan, B. and Leng, P. (2009). The role of ABA in triggering ethylene biosynthesis and ripening of tomato fruit. Journal of Experimental Botany. 60: 1579–1588.
Zhao, D.Y., Shen, L., Fan, B., Liu, K.L., Yu, M.M., Zheng, Y., Ding, Y. and Sheng, J.P. (2009). Physiological and genetic properties of tomato fruits from 2 cultivars differing in chilling tolerance at cold storage. Food Chemistry. 74: 348–352.
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Asadi, S., Moghaddam, M., Ghasemi pirbalouti, A. and Fotovat, A. (2018). Evaluation of physiological characteristics and antioxidant activity of sweet basil (Ocimum basilicum cv. Keshkeni luvelou) under different levels of methyl jasmonate and lead toxicity. Journal of Iranian Plant Ecophysiological Research. 13(51): 1-16.
Alscher, R.G. and Cumming, J.R. (1990). Stress responses in plants: adaptation and acclimation Mechanisms. John Wiley and Sons. INC Publication.
Banerjee, S., Yuan, X., Germida, J.J. and Vujanovic V. (2014). Gene expression patterns in wheat coleorhiza under cold-and biological stratification. Microbiological research. 169(7-8): 616-622.
Bose, J., Rodrigo-Moreno A. and Shabala, S. (2014). ROS homeostasis in halophytes in the context of salinity tolerance. Journal of Experimental Botany. 65: 1241–1257.
Brand-Williams, W., Cuvelier, M.E. andBerset, C. (1995). Use of free radical method to evaluate antioxidant activity. LWT- Food Science and Technology. 28(1):25-30
Bunn, R., Lekberg, Y. and Zabinski, C. (2009). Arbuscular mycorrhizal fungi ameliorate temperature stress in thermophilic plants. Ecology. 90:1378–1388.
Davies, P.J. and Mansfield, TA. (1983). The role of abscisic acid in drought avoidance, In: Abscisic acid (ed. by F.T. Adicott). Praeger, New York, 237-268.
Elhamirad, A.H. and Zamanipoor, M.H. (2012). Thermal stability of some flavonoids and phenolic acids in sheep tallow olein. European Journal of Lipid Science and Technology. 114(5): 602-606.
Ghorbanli, M., Rostami Abousaeidi, M. and Bakhshi Khaniki, Gh.R. (2014). Effect of naphthalene acetic acid (NAA) and abscisic acid (ABA) on quality and quantity of Strawberry Selva cultivar (Fragaria ananassa cv. Selva) and Paros cultivar (Fragaria ananassa cv. Paros). Journal of Iranian Plant Ecophysiological Research. 9: 30-39.
Großkinsky, D.K., Naseem, M., Abdelmohsen, U.R., Plickert, N., Engelke. T., Griebel, T., Zeier. J., Novák, O., Strnad, M., Pfeifhofer, H., Graaff, E.V.D. Simon, U. and Roitsch, T. (2011). Cytokinins mediate resistance against Pseudomonas syringaein tobacco through increased antimicrobial phytoalexin synthesis independent of salicylic acid signaling. Plant Physiol, 157:815–830
Hiwasa, K., Kinugasa, Y., Amano, S., Hashimoto, A., Nakano, R., Inaba, A. and Kubo, Y. (2003). Ethylene is required for both the initiation and progression of softening in pear Pyrus communis L. fruit. Journal of Experimental Botany. 54: 771–779.
Hosseinzad, B.E., Chaparzadeh N. and Dilmaghani K. (2014). Effect of salicylic acid on growth parameters, osmolytes and osmotic potential in radish (Raphanus sativus L.) under salt stress. Journal of Plant Research. 27(1): 32-40. (In Persian)
Huber, D.J. (1983). Polyuronide degradation and hemicellulose modification on ripening tomato fruit. Journal of the American Society for Horticultural Science. 108: 405–409.
Jahan tigh haghighi, Z., Fahmideh, L., Fazeli nasab, B. (2018). Evaluation and comparison of Leaf antioxidant properties and morphological traits of tomato varieties (Lycopersicon esculentum L). 13(50): 63-76.
Jajic, I., Sarna, T. and Strzalka, K. (2015). Senescence, Stress, and Reactive Oxygen Species. Plants. 4(3): 393-411.
Jamshidi, M., Ahmadi, H.R., Rezazadeh, Sh., Fathi, F. and Mazanderani, M. (2010). Study on phenolicd and anioxidant activity of some selected plant of Mazandaran province. Medic Plan. 934:177-183. (In Persian).
Ji, K., Kai, W., Zhao, B., Sun, Y. and Yuan, B. (2014). SlNCED1and SlCYP707A2: key genes involved in ABA metabolism during tomato fruit ripening. Journal of Experimental Botany. 6518: 5243-5255.
Kilian, J., Whitehead, D., Horak, J., Wanke, D., Weinl, S., Batistic, O.D., Angelo, C., Bornberg-Bauer, E., Kudla, J. and Harter, K. (2007). The AtGen Express global stress expression data set: protocols, evolution and model data analysis of UV-B light, drought and cold stress responses. The Plant Journal. 50:347–363.
Kok, E., Islek, C. and Sulun Ustun A. (2010). Effect of Cold on Protein, Proline, Phenolic Compounds and Chlorophyll Content of Two Pepper Capsicum annuum L. Varieties. G.U. Journal of Science. 231: 1-6.
Leon, P. and Sheen, J. (2003). Sugar and hormone connections. Trends Plant Science. 8: 110-116.
Liang, L.H., Mei, X., Lin, F., Xia, J., Liu, S.J. and Wang, J.H. (2009). Effect of low temperature stress on tissue structure and physiological index of cashew young leaves. Journal of Ecology and Environmental Sciences. 18: 317-320.
Mohammadi, B., Zia Jahromi, N., Sadeghi, H. and Mirzaei, A. (2016). Compare the amount of phenols, flavonoids and antioxidant activity of five varieties of Iranian olive leaf hydroalcoholic extract. Bringing Knowledge, 20 (10): 888-898. (In Persian).
Munder, M. (2009). Arginase: an emerging key player in the mammalian immune system: review. British Journal of Pharmacology. 158: 638–651.
Nakamura, S., Chono, M., Abe, F. and Miura, H. (2010). Mapping a diploid wheat abscisic acid 8′-hydroxylase homologue in the seed dormancy QTL region on chromosome 5Am. Euphytica. 171(1):111.
Oliver, S.N., Dennis, E.S. and Dolferus, R. (2007). ABA regulates apoplastic sugar transport and is a potential signal for cold-induced pollen sterility in rice. Plant and Cell physiology. 48(9): 1319-1330.
Pakkish, Z., Rahemi, M. and Baghizadeh, A. (2009). Seasonal changes of peroxidase, polyphenol oxidase enzyme activity and phenol content during and after rest in pistachio Pistacia vera L. Flower Buds. World Applied Sciences. 69: 1193-1199.
Qin, X.Q. and Zeevaart J.A. (1999). The 9-cis-epoxycarotenoid cleavage reaction is the key regulatory step of abscisic acid biosynthesis in water-stressed bean. Proceedings of the National Academy of sciences. 96(26): 15354-15361.
Qin, X.Q. and Zeevaart, J.A.D. (2002). Overexpression of a 9-cis-epoxycarotenoid dioxygenase gene in Nicotiana plumbaginifolia increases abscisic acid and phasic acid levels and enhances drought tolerance. Plant Physiology. 128: 544–551.
Raghavendra, A.S., Gonugunta, V.K., Christmann, A. and Grill, E. (2010). ABA perception and signaling. Trends Plant Science. 15:395–401.
Rivero, R.M., Ruiz, J.M., Garcıa, P.C., Lopez-Lefebre, L.R., Sanchez, E. and Romero, L. (2001). Resistance to cold and heat stress: accumulation of phenolic compounds in tomato and watermelon plants. Plant science. 160: 315-321.
Salmanian, S., Sadeghi Mahoonak, A. Alami, M. and Ghorbani, M. (2014). Evaluation of Total Phenolic, Flavonoid, Anthocyanin Compounds, Antibacterial and Antioxidant Activity of Hawthorn (Crataegus Elbursensis) Fruit Acetonic Extract. Journal of Rafsanjan University of Medical Sciences. 13 (1): 53-66. (In Persian)
Sangwan, V. and Dhindsa, R.S. (2002). In vivo and in vitro activation of temperature-responsive plant map kinases. FEBS Letters. 531:561–564.
Sayyari, M., Babalar, M., Kalantari, S., MartinezRomero, D., Guillen, F., Serrano, M. and Valero, D. (2011). Vapour treatments with methyl salicylate or methyl jasmonate alleviated chilling injury and enhanced antioxidant potential during postharvest storage of pomegranates. Food Chemistry. 124: 964–970.
Schlegl, Z. (1996). Photo assimilate distribution in plants and crops, source-sink relationships. Environmental Control in Biology. 323-330.
Schwartz, S.H., Qin, X.Q. and Zeevaart, J.A.D. (2003). Elucidation of the indirect pathway of abscisic acid biosynthesis by mutants, genes, and enzymes. Plant Physiology. 131:1591–1601.
Seppanen, M.M. (2000). Characterize of freezing tolerance in Solanum commersoniidun. with special reference of the relationship between and oxidative stress. University of Helsinki department of production section of crop husbandry. 56:4-44.
Sun, L., Wang, Y.P., Chen, P., Ren, J., Ji, K., Li, Q., Li, P., Dai, S.J. and Leng, P. (2011). Transcriptional regulation of SlPYL, SlPP2C, and SlSnRK2 gene families encoding ABA signal core components during tomato fruit development and drought stress. Journal of Experimental Botany. 62:5659–5669.
Taffazoli, A.A., and Kalabil, R. (2002). Increasing sugars and cold resistance in Kiwi fruit by ABA hormone. Iran Agricultural Research. 11. (In Persian)
Taylor, I.B., Sonneveld, T., Bugg, T.D.H. and Thompson, A.J. (2005). Regulation and manipulation of the biosynthesis of abscisic acid, including the supply of xanthophyll precursors. Journal of Plant Growth Regulation. 24:253–273.
Van kiet, H., Nose, A. and Shao, H. (2016). Effect of Cold Stress on Root Growth, Accumulation of Soluble Proteins and Free Amino Acids of Sheath Blight-Resistant Rice Genotype 32R. Tropical Agriculture and Development. 60(3): 191-194.
Wang, G.D., Liu, Q., Shang, X.T., Chen, C., Xu, N., Guan, J. and Meng, Q.T. (2017). Overexpression of transcription factor SlNAC35 enhances the chilling tolerance of transgenic tomato. Biologia Plantarum. 62(3):479–488.
Weidner, S., Kordala, E., Brosowska-Arendt, W., Karamac, M., Kosinska, A. and Amarowicz, R. (2009). Phenolic compounds and properties of antioxidants in grapevine roots Vitis vinifera L. under low temperature stress followed by recovery. Acta Societatis Botanicorum Poloniae. 78: 279-286.
Went, F.W. (1953). The effect of temperature on plant growth. Annual Review of Plant Physiology. 4:347–362.
Xiong, L., Wangm R.G., Mao, G. and Koczan, J.M. (2006). Identification of drought tolerance determinants by genetic analysis of root response to drought stress and abscisic acid. Plant Physiology. 142: 1065–1074.
Zhang, M., Yuan, B. and Leng, P. (2009). The role of ABA in triggering ethylene biosynthesis and ripening of tomato fruit. Journal of Experimental Botany. 60: 1579–1588.
Zhao, D.Y., Shen, L., Fan, B., Liu, K.L., Yu, M.M., Zheng, Y., Ding, Y. and Sheng, J.P. (2009). Physiological and genetic properties of tomato fruits from 2 cultivars differing in chilling tolerance at cold storage. Food Chemistry. 74: 348–352.