تعیین کلون‌های متحمل به تنش خشکی چای (Camellia sinensis L.) با بررسی فعالیت آنزیم‌های آنتیاکسیدانت
الموضوعات :
مهدی رحیمی
1
,
مجتبی کردرستمی
2
,
مجتبی مرتضوی
3
,
صنم صفایی چایی کار
4
1 - گروه بیوتکنولوژی، پژوهشکده علوم محیطی، پژوهشگاه علوم و تکنولوژی پیشرفته و علوم محیطی،
دانشگاه تحصیلات تکمیلی صنعتی و فناوری پیشرفته، کرمان، ایران.
2 - گروه بیوتکنولوژی، دانشکده علوم کشاورزی، دانشگاه گیلان، رشت، ایران.
3 - گروه بیوتکنولوژی، پژوهشکده علوم محیطی، پژوهشگاه علوم و تکنولوژی پیشرفته و علوم محیطی،
دانشگاه تحصیلات تکمیلی صنعتی و فناوری پیشرفته، کرمان، ایران.
4 - پژوهشکده چای، موسسه تحقیقات علوم باغبانی، سازمان تحقیقات، آموزش و ترویج کشاورزی، لاهیجان، ایران.
الکلمات المفتاحية: پراکسیداز, کاتالاز, تحمل به خشکی, بتاکاروتن, مالون‌دیآلدهید,
ملخص المقالة :
به‌منظور بررسی فعالیت آنزیم‌های آنتیاکسیدانت 14 کلون چای در شرایط نرمال و تنش خشکی، دو آزمایش جداگانه به‌صورت طرح بلوک‌های کامل تصادفی با دو تکرار در سال زراعی 1396 در ایستگاه تحقیقات چای فشالم (رشت) اجرا گردید. در هر دو طرح، آبیاری به‌صورت معمول تا آخر تیر ماه انجام شد؛ اما در تیمار خشکی، از ابتدای مرداد تا موقع برداشت برگ‌های چای، آبیاری قطع گردید. در آخر مرداد، از هر کرت در هر دو آزمایش، برگ کلون‌های چای برداشته شده و به فریزر 80- منتقل شدند. فعالیت آنزیم‌های آنتیاکسیدانت آسکوربات پراکسیداز، سوپراُکسیددیس‌موتاز، کاتالاز، فنیل‌آلانین آمونیالیاز، لیپید پراکسیداز، میزان مالون‌دی‌آلدئید، بتاکاروتن و لیکوپن اندازه‌گیری شدند. نتایج نشان داد که تنش خشکی باعث افزایش فعالیت آنزیم‌های آنتیاکسیدانت گردید. بر اساس نتایج به‌دست‌آمده از مقایسات میانگین، کلون‌های 100، 399 و بذری تحت شرایط تنش خشکی دارای بیشترین مقادیر برای بیشترین مقدار فعالیت آنزیم‌های آنتیاکسیدانت آسکوربات پراکسیداز، سوپراُکسیددیس‌موتاز، کاتالاز، پراکسیداز، فنیل‌آلانین آمونیالیاز، لیپید پراکسیداز بودند. در طرف مقابل، این کلو‌ن‌ها کمترین مقدار را برای مالون‌دی‌آلدئید دارا بودند. این مساله تحمل نسبی آن‌ها را به خشکی نشان می‌دهد. کلون‌های 278 و 276 با کمترین مقدار آنزیم‌های آنتی اکسیدانت در شرایط تنش خشکی (بجز مالون‌دی‌آلدئید) به‌عنوان کلون‌های حساس در نظر گرفته شدند.
Alscher, R.G., Erturk, N. and Heath, L.S. (2002). Role of superoxide dismutases (SODs) in controlling oxidative stress in plants. Journal of Experimental Botany. 53: 1331-1341.
Apel, K. and Hirt, H. (2004). Reactive oxygen species: metabolism, oxidative stress, and signal transduction. Annual Review of Plant Biology. 55: 373-399.
Ashraf, M. (2009). Biotechnological approach of improving plant salt tolerance using antioxidants as markers. Biotechnology Advances. 27: 84-93.
Ashraf, M. and Harris, P. (2004). Potential biochemical indicators of salinity tolerance in plants. Plant Science. 166: 3-16.
Badawi, G.H., Kawano, N., Yamauchi, Y., Shimada, E., Sasaki, R., Kubo, A. and Tanaka, K. (2004). Over‐expression of ascorbate peroxidase in tobacco chloroplasts enhances the tolerance to salt stress and water deficit. Physiologia Plantarum. 121: 231-238.
Beauchamp, C. and Fridovich, I. (1971). Superoxide dismutase: improved assays and an assay applicable to acrylamide gels. Analytical Biochemistry. 44: 276-287.
Beers, R.F. and Sizer, I.W. (1952). A spectrophotometric method for measuring the breakdown of hydrogen peroxide by catalase. The Journal of Biological Chemistry. 195: 133-140.
Borsani, O., Valpuesta, V. and Botella, M.A. (2001). Evidence for a role of salicylic acid in the oxidative damage generated by NaCl and osmotic stress in Arabidopsis seedlings. Plant Physiology. 126: 1024-1030.
Bowler, C., Montagu, M.v. and Inzé, D. (1992). Superoxide dismutase and stress tolerance. Annual Review of Plant Biology. 43: 83-116.
Bowler, C., Van Camp, W., Van Montagu, M., Inze, D. and Asada, K. (1994). Superoxide dismutase in plants. Critical Reviews in Plant Sciences. 13: 199-218.
Cai‐Hong, P., Su‐Jun, Z., Zhi‐Zhong, G. and Bao‐Shan, W. (2005). NaCl treatment markedly enhances H2O2‐scavenging system in leaves of halophyte Suaeda salsa. Physiologia Plantarum. 125: 490-499.
Chance, B. and Maehly, A. (1955). Assay of catalases and peroxidases. Academic Press, New York.
Ciarmiello, L.F., Woodrow, P., Fuggi, A., Pontecorvo, G. and Carillo, P. (2011). Plant genes for abiotic stress. In: Shanker, A. and Venkateswarlu , B. (Eds.), Abiotic stress in plants- mechanisms and adaptations. InTech, London, United Kingdom, pp. 284-308.
Cicek, N. and Cakirlar, H. (2008). Changes in some antioxidant enzyme activities in six soybean cultivars in response to long-term salinity at two different temperatures. General and Applied Plant Physiology. 34: 267-280.
Cornic, G., Ghashghaie, J., Genty, B. and Briantais, J. (1992). Leaf photosynthesis is resistant to a mild drought stress. Photosynthetica. 27: 295-309.
Cross, A.R., Erickson, R.W., Ellis, B.A. and Curnutte, J.T. (1999). Spontaneous activation of NADPH oxidase in a cell-free system: unexpected multiple effects of magnesium ion concentrations. Biochemical Journal. 338: 229-233.
Demiral, T. and Türkan, I. (2005). Comparative lipid peroxidation, antioxidant defense systems and proline content in roots of two rice cultivars differing in salt tolerance. Environmental and Experimental Botany. 53: 247-257.
Dufossé, L., Galaup, P., Yaron, A., Arad, S.M., Blanc, P., Murthy, K.N.C. and Ravishankar, G.A. (2005). Microorganisms and microalgae as sources of pigments for food use: a scientific oddity or an industrial reality? Trends in Food Science & Technology. 16: 389-406.
FAOSTAT (2012). Food and Agriculture Organization of the United Nations, FAOSTAT database, available at http://faostat3.fao.org/faostat-gateway/go/to/download/Q/QC/E.
Foyer, C.H. and Noctor, G. (2005). Oxidant and antioxidant signalling in plants: a re‐evaluation of the concept of oxidative stress in a physiological context. Plant, Cell & Environment. 28: 1056-1071.
Gossett, D.R., Millhollon, E.P. and Lucas, M. (1994). Antioxidant response to NaCl stress in salt-tolerant and salt-sensitive cultivars of cotton. Crop Science. 34: 706-714.
Halliwell, B. (1993). The chemistry of free radicals. Toxicology and Industrial Health. 9: 1-21.
Haupt‐Herting, S. and Fock, H.P. (2000). Exchange of oxygen and its role in energy dissipation during drought stress in tomato plants. Physiologia Plantarum. 110: 489-495.
Heath, R.L. and Packer, L. (1968). Photoperoxidation in isolated chloroplasts: I. Kinetics and stoichiometry of fatty acid peroxidation. Archives of Biochemistry and Biophysics. 125: 189-198.
Hernández, J.A., Ferrer, M.A., Jiménez, A., Barceló, A.R. and Sevilla, F. (2001). Antioxidant Systems and O2.−/H2O2 production in the apoplast of pea leaves. Its relation with salt-induced necrotic lesions in minor veins. Plant Physiology. 127: 817-831.
Huseynova, I.M., Suleymanov, S.Y. and Rustamova, S.M. (2010). Response of photosynthetic apparatus and antioxidant defense systems in Triticum aestivum L. genotypes subjected to drought stress. Proceedings of ANAS (Biological Sciences). 65: 49-59.
Kjalke, M., Andersen, M.B., Schneider, P., Christensen, B., Schülein, M. and Welinder, K.G. (1992). Comparison of structure and activities of peroxidases from Coprinus cinereus, Coprinus macrorhizus and Arthromyces ramosus. Biochimica et Biophysica Acta (BBA)-Protein Structure and Molecular Enzymology. 1120: 248-256.
Lin, C.C. and Kao, C.H. (2001). Cell wall peroxidase activity, hydrogen peroxide level and NaCl-inhibited root growth of rice seedlings. Plant and Soil. 230: 135-143.
Manavalan, L.P., Guttikonda, S.K., Phan Tran, L.-S. and Nguyen, H.T. (2009). Physiological and molecular approaches to improve drought resistance in soybean. Plant and Cell Physiology. 50: 1260-1276.
Masoudian, Z., Norastehnia, A. and Falakroo, K. (2014). Study of drought tolerance in selective clones of tea (Camellia sinensis L.). Iranian Journal of Plant Biology. 6: 155-170.
Mazel, A., Leshem, Y., Tiwari, B.S. and Levine, A. (2004). Induction of salt and osmotic stress tolerance by overexpression of an intracellular vesicle trafficking protein AtRab7 (AtRabG3e). Plant Physiology. 134: 118-128.
Meloni, D.A., Oliva, M.A., Martinez, C.A. and Cambraia, J. (2003). Photosynthesis and activity of superoxide dismutase, peroxidase and glutathione reductase in cotton under salt stress. Environmental and Experimental Botany. 49: 69-76.
Mhamdi, A., Queval, G., Chaouch, S., Vanderauwera, S., Van Breusegem, F. and Noctor, G. (2010). Catalase function in plants: a focus on Arabidopsis mutants as stress-mimic models. Journal of Experimental Botany. 61: 4197-4220.
Mittler, R. (2002). Oxidative stress, antioxidants and stress tolerance. Trends in Plant Science. 7: 405-410.
Nagata, M. and Yamashita, I. (1992). Simple method for simultaneous determination of chlorophyll and carotenoids in tomato fruit. Nippon Shokuhin Kogyo Gakkaishi. 39: 925-928.
Nakano, Y. and Asada, K. (1981). Hydrogen peroxide is scavenged by ascorbate-specific peroxidase in spinach chloroplasts. Plant and Cell Physiology. 22: 867-880.
Nogués, S. and Baker, N.R. (2000). Effects of drought on photosynthesis in Mediterranean plants grown under enhanced UV‐B radiation. Journal of Experimental Botany. 51: 1309-1317.
Omidi, H. (2010). Changes of proline content and activity of antioxidative enzymes in two canola genotype under drought stress. American Journal of Plant Physiology. 5: 338-349.
Parida, A.K. and Das, A.B. (2005). Salt tolerance and salinity effects on plants: a review. Ecotoxicology and Environmental Safety. 60: 324-349.
Reddy, A.R., Chaitanya, K.V. and Vivekanandan, M. (2004). Drought-induced responses of photosynthesis and antioxidant metabolism in higher plants. Journal of Plant Physiology. 161: 1189-1202.
Sekmen, A.H., Türkan, İ. and Takio, S. (2007). Differential responses of antioxidative enzymes and lipid peroxidation to salt stress in salt‐tolerant Plantago maritima and salt‐sensitive Plantago media. Physiologia Plantarum. 131: 399-411.
Seraji, A., Pourjam, E., Tanha, M.Z. and Safaei, N. (2007). Biology and population dynamics of tea root lesion nematode (Pratylenchus loosi) in Iran. Iranian Journal of Plant Pathology. 43: 98-115. [In Persian with English Summary].
Sharma, P., Jha, A.B., Dubey, R.S. and Pessarakli, M. (2012). Reactive oxygen species, oxidative damage, and antioxidative defense mechanism in plants under stressful conditions. Journal of Botany. 2012.
Ślesak, I., Miszalski, Z., Karpinska, B., Niewiadomska, E., Ratajczak, R. and Karpinski, S. (2002). Redox control of oxidative stress responses in the C3–CAM intermediate plant Mesembryanthemum crystallinum. Plant Physiology and Biochemistry. 40: 669-677.
Stepien, P. and Klobus, G. (2005). Antioxidant defense in the leaves of C3 and C4 plants under salinity stress. Physiologia Plantarum. 125: 31-40.
Sudhakar, C., Lakshmi, A. and Giridarakumar, S. (2001). Changes in the antioxidant enzyme efficacy in two high yielding genotypes of mulberry (Morus alba L.) under NaCl salinity. Plant Science. 161: 613-619.
Tester, M. and Bacic, A. (2005). Abiotic stress tolerance in grasses. From model plants to crop plants. Plant Physiology. 137: 791–793.
Van Breusegem, F. and Dat, J.F. (2006). Reactive oxygen species in plant cell death. Plant Physiology. 141: 384-390.
Wang, J.W., Zheng, L.P., Wu, J.Y. and Tan, R.X. (2006). Involvement of nitric oxide in oxidative burst, phenylalanine ammonia-lyase activation and Taxol production induced by low-energy ultrasound in Taxus yunnanensis cell suspension cultures. Nitric Oxide. 15: 351-358.
Wang, W.-B., Kim, Y.-H., Lee, H.-S., Kim, K.-Y., Deng, X.-P. and Kwak, S.-S. (2009). Analysis of antioxidant enzyme activity during germination of alfalfa under salt and drought stresses. Plant Physiology and Biochemistry. 47: 570-577.
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Alscher, R.G., Erturk, N. and Heath, L.S. (2002). Role of superoxide dismutases (SODs) in controlling oxidative stress in plants. Journal of Experimental Botany. 53: 1331-1341.
Apel, K. and Hirt, H. (2004). Reactive oxygen species: metabolism, oxidative stress, and signal transduction. Annual Review of Plant Biology. 55: 373-399.
Ashraf, M. (2009). Biotechnological approach of improving plant salt tolerance using antioxidants as markers. Biotechnology Advances. 27: 84-93.
Ashraf, M. and Harris, P. (2004). Potential biochemical indicators of salinity tolerance in plants. Plant Science. 166: 3-16.
Badawi, G.H., Kawano, N., Yamauchi, Y., Shimada, E., Sasaki, R., Kubo, A. and Tanaka, K. (2004). Over‐expression of ascorbate peroxidase in tobacco chloroplasts enhances the tolerance to salt stress and water deficit. Physiologia Plantarum. 121: 231-238.
Beauchamp, C. and Fridovich, I. (1971). Superoxide dismutase: improved assays and an assay applicable to acrylamide gels. Analytical Biochemistry. 44: 276-287.
Beers, R.F. and Sizer, I.W. (1952). A spectrophotometric method for measuring the breakdown of hydrogen peroxide by catalase. The Journal of Biological Chemistry. 195: 133-140.
Borsani, O., Valpuesta, V. and Botella, M.A. (2001). Evidence for a role of salicylic acid in the oxidative damage generated by NaCl and osmotic stress in Arabidopsis seedlings. Plant Physiology. 126: 1024-1030.
Bowler, C., Montagu, M.v. and Inzé, D. (1992). Superoxide dismutase and stress tolerance. Annual Review of Plant Biology. 43: 83-116.
Bowler, C., Van Camp, W., Van Montagu, M., Inze, D. and Asada, K. (1994). Superoxide dismutase in plants. Critical Reviews in Plant Sciences. 13: 199-218.
Cai‐Hong, P., Su‐Jun, Z., Zhi‐Zhong, G. and Bao‐Shan, W. (2005). NaCl treatment markedly enhances H2O2‐scavenging system in leaves of halophyte Suaeda salsa. Physiologia Plantarum. 125: 490-499.
Chance, B. and Maehly, A. (1955). Assay of catalases and peroxidases. Academic Press, New York.
Ciarmiello, L.F., Woodrow, P., Fuggi, A., Pontecorvo, G. and Carillo, P. (2011). Plant genes for abiotic stress. In: Shanker, A. and Venkateswarlu , B. (Eds.), Abiotic stress in plants- mechanisms and adaptations. InTech, London, United Kingdom, pp. 284-308.
Cicek, N. and Cakirlar, H. (2008). Changes in some antioxidant enzyme activities in six soybean cultivars in response to long-term salinity at two different temperatures. General and Applied Plant Physiology. 34: 267-280.
Cornic, G., Ghashghaie, J., Genty, B. and Briantais, J. (1992). Leaf photosynthesis is resistant to a mild drought stress. Photosynthetica. 27: 295-309.
Cross, A.R., Erickson, R.W., Ellis, B.A. and Curnutte, J.T. (1999). Spontaneous activation of NADPH oxidase in a cell-free system: unexpected multiple effects of magnesium ion concentrations. Biochemical Journal. 338: 229-233.
Demiral, T. and Türkan, I. (2005). Comparative lipid peroxidation, antioxidant defense systems and proline content in roots of two rice cultivars differing in salt tolerance. Environmental and Experimental Botany. 53: 247-257.
Dufossé, L., Galaup, P., Yaron, A., Arad, S.M., Blanc, P., Murthy, K.N.C. and Ravishankar, G.A. (2005). Microorganisms and microalgae as sources of pigments for food use: a scientific oddity or an industrial reality? Trends in Food Science & Technology. 16: 389-406.
FAOSTAT (2012). Food and Agriculture Organization of the United Nations, FAOSTAT database, available at http://faostat3.fao.org/faostat-gateway/go/to/download/Q/QC/E.
Foyer, C.H. and Noctor, G. (2005). Oxidant and antioxidant signalling in plants: a re‐evaluation of the concept of oxidative stress in a physiological context. Plant, Cell & Environment. 28: 1056-1071.
Gossett, D.R., Millhollon, E.P. and Lucas, M. (1994). Antioxidant response to NaCl stress in salt-tolerant and salt-sensitive cultivars of cotton. Crop Science. 34: 706-714.
Halliwell, B. (1993). The chemistry of free radicals. Toxicology and Industrial Health. 9: 1-21.
Haupt‐Herting, S. and Fock, H.P. (2000). Exchange of oxygen and its role in energy dissipation during drought stress in tomato plants. Physiologia Plantarum. 110: 489-495.
Heath, R.L. and Packer, L. (1968). Photoperoxidation in isolated chloroplasts: I. Kinetics and stoichiometry of fatty acid peroxidation. Archives of Biochemistry and Biophysics. 125: 189-198.
Hernández, J.A., Ferrer, M.A., Jiménez, A., Barceló, A.R. and Sevilla, F. (2001). Antioxidant Systems and O2.−/H2O2 production in the apoplast of pea leaves. Its relation with salt-induced necrotic lesions in minor veins. Plant Physiology. 127: 817-831.
Huseynova, I.M., Suleymanov, S.Y. and Rustamova, S.M. (2010). Response of photosynthetic apparatus and antioxidant defense systems in Triticum aestivum L. genotypes subjected to drought stress. Proceedings of ANAS (Biological Sciences). 65: 49-59.
Kjalke, M., Andersen, M.B., Schneider, P., Christensen, B., Schülein, M. and Welinder, K.G. (1992). Comparison of structure and activities of peroxidases from Coprinus cinereus, Coprinus macrorhizus and Arthromyces ramosus. Biochimica et Biophysica Acta (BBA)-Protein Structure and Molecular Enzymology. 1120: 248-256.
Lin, C.C. and Kao, C.H. (2001). Cell wall peroxidase activity, hydrogen peroxide level and NaCl-inhibited root growth of rice seedlings. Plant and Soil. 230: 135-143.
Manavalan, L.P., Guttikonda, S.K., Phan Tran, L.-S. and Nguyen, H.T. (2009). Physiological and molecular approaches to improve drought resistance in soybean. Plant and Cell Physiology. 50: 1260-1276.
Masoudian, Z., Norastehnia, A. and Falakroo, K. (2014). Study of drought tolerance in selective clones of tea (Camellia sinensis L.). Iranian Journal of Plant Biology. 6: 155-170.
Mazel, A., Leshem, Y., Tiwari, B.S. and Levine, A. (2004). Induction of salt and osmotic stress tolerance by overexpression of an intracellular vesicle trafficking protein AtRab7 (AtRabG3e). Plant Physiology. 134: 118-128.
Meloni, D.A., Oliva, M.A., Martinez, C.A. and Cambraia, J. (2003). Photosynthesis and activity of superoxide dismutase, peroxidase and glutathione reductase in cotton under salt stress. Environmental and Experimental Botany. 49: 69-76.
Mhamdi, A., Queval, G., Chaouch, S., Vanderauwera, S., Van Breusegem, F. and Noctor, G. (2010). Catalase function in plants: a focus on Arabidopsis mutants as stress-mimic models. Journal of Experimental Botany. 61: 4197-4220.
Mittler, R. (2002). Oxidative stress, antioxidants and stress tolerance. Trends in Plant Science. 7: 405-410.
Nagata, M. and Yamashita, I. (1992). Simple method for simultaneous determination of chlorophyll and carotenoids in tomato fruit. Nippon Shokuhin Kogyo Gakkaishi. 39: 925-928.
Nakano, Y. and Asada, K. (1981). Hydrogen peroxide is scavenged by ascorbate-specific peroxidase in spinach chloroplasts. Plant and Cell Physiology. 22: 867-880.
Nogués, S. and Baker, N.R. (2000). Effects of drought on photosynthesis in Mediterranean plants grown under enhanced UV‐B radiation. Journal of Experimental Botany. 51: 1309-1317.
Omidi, H. (2010). Changes of proline content and activity of antioxidative enzymes in two canola genotype under drought stress. American Journal of Plant Physiology. 5: 338-349.
Parida, A.K. and Das, A.B. (2005). Salt tolerance and salinity effects on plants: a review. Ecotoxicology and Environmental Safety. 60: 324-349.
Reddy, A.R., Chaitanya, K.V. and Vivekanandan, M. (2004). Drought-induced responses of photosynthesis and antioxidant metabolism in higher plants. Journal of Plant Physiology. 161: 1189-1202.
Sekmen, A.H., Türkan, İ. and Takio, S. (2007). Differential responses of antioxidative enzymes and lipid peroxidation to salt stress in salt‐tolerant Plantago maritima and salt‐sensitive Plantago media. Physiologia Plantarum. 131: 399-411.
Seraji, A., Pourjam, E., Tanha, M.Z. and Safaei, N. (2007). Biology and population dynamics of tea root lesion nematode (Pratylenchus loosi) in Iran. Iranian Journal of Plant Pathology. 43: 98-115. [In Persian with English Summary].
Sharma, P., Jha, A.B., Dubey, R.S. and Pessarakli, M. (2012). Reactive oxygen species, oxidative damage, and antioxidative defense mechanism in plants under stressful conditions. Journal of Botany. 2012.
Ślesak, I., Miszalski, Z., Karpinska, B., Niewiadomska, E., Ratajczak, R. and Karpinski, S. (2002). Redox control of oxidative stress responses in the C3–CAM intermediate plant Mesembryanthemum crystallinum. Plant Physiology and Biochemistry. 40: 669-677.
Stepien, P. and Klobus, G. (2005). Antioxidant defense in the leaves of C3 and C4 plants under salinity stress. Physiologia Plantarum. 125: 31-40.
Sudhakar, C., Lakshmi, A. and Giridarakumar, S. (2001). Changes in the antioxidant enzyme efficacy in two high yielding genotypes of mulberry (Morus alba L.) under NaCl salinity. Plant Science. 161: 613-619.
Tester, M. and Bacic, A. (2005). Abiotic stress tolerance in grasses. From model plants to crop plants. Plant Physiology. 137: 791–793.
Van Breusegem, F. and Dat, J.F. (2006). Reactive oxygen species in plant cell death. Plant Physiology. 141: 384-390.
Wang, J.W., Zheng, L.P., Wu, J.Y. and Tan, R.X. (2006). Involvement of nitric oxide in oxidative burst, phenylalanine ammonia-lyase activation and Taxol production induced by low-energy ultrasound in Taxus yunnanensis cell suspension cultures. Nitric Oxide. 15: 351-358.
Wang, W.-B., Kim, Y.-H., Lee, H.-S., Kim, K.-Y., Deng, X.-P. and Kwak, S.-S. (2009). Analysis of antioxidant enzyme activity during germination of alfalfa under salt and drought stresses. Plant Physiology and Biochemistry. 47: 570-577.
