بررسی ویژگیهای فتوسنتزی و فیتوشیمیایی برخی از اکوتیپهای بنفشه معطر (Viola odorata L.) در استانهای مازندران و گلستان
محورهای موضوعی : ژنتیکحسین مرادی 1 , مهدی حدادی نژاد 2 , علیرضا یاوری 3 , مهدی محمدی ازنی 4 , سید مریم موسوی 5 , سیدمحمدامین حسینی 6
1 - گروه علوم و مهندسی باغبانی، دانشکده علوم زراعی و پژوهشکده فناوریهای زیستی گیاهان دارویی
دانشگاه علوم کشاورزی و منابع طبیعی ساری، ایران
2 - گروه علوم و مهندسی باغبانی، دانشکده علوم زراعی، دانشگاه علوم کشاورزی و منابع طبیعی ساری، ایران
3 - استادیار گروه علوم و مهندسی باغبانی، دانشکده کشاورزی و منابع طبیعی، دانشگاه هرمزگان، بندرعباس، ایران
4 - گروه علوم و مهندسی باغبانی، دانشکده علوم زراعی، دانشگاه علوم کشاورزی و منابع طبیعی ساری، ایران
5 - گروه علوم و مهندسی باغبانی، دانشکده علوم زراعی، دانشگاه علوم کشاورزی و منابع طبیعی ساری، ایران
6 - گروه علوم و مهندسی باغبانی، دانشکده علوم زراعی، دانشگاه علوم کشاورزی و منابع طبیعی ساری، ایران
کلید واژه: اکوتیپ, صفات مورفولوژیکی, فلاونوئید, ترکیبات فیتوشیمیایی, بنفشه ایرانی,
چکیده مقاله :
به منظور ارزیابی تنوع ژنتیکی با استفاده از ویژگی های ظاهری و فیتوشیمیایی گیاه بنفشه معطر (Viola odorata L.)، 10 منطقه از استان های مازندران و گلستان بررسی شد. صفات فیتوشیمیایی نظیر میزان ترکیبات فنلی، فلاونوئیدها، ظرفیت آنتی اکسیدانی، کلروفیل a، کلروفیل b، کلروفیل کل و صفات ریخت شناسی از جمله طول و عرض برگ، طول دم برگ و طول ریشه ارزیابی گردید. در بررسی صفات فیتوشیمیایی ژنوتیپ های مناطق ساری و کسوت دارای بالاترین میزان کلروفیل b و کلروفیل کل بوده و اکوتیب ساری از بالاترین میزان فلاونوئید بر خوردار بودند. از نظر میزان ظرفیت آنتی اکسیدانی، دو اکوتیپ ساری و وری، با غلظت مهار 60 درصد رادیکال های آزاد بالاترین مقدار را نشان دادند. اکوتیب های مناطق زنگلاب، آق چشمه، تنگه چهل چای با ارتفاع از سطح دریای کمتر نسبت به مناطق دیگر مورد مطالعه، بالاترین میزان فنل کل را به میزان 25 میلیگرم اسید گالیک در گرم وزن خشک را نشان دادند. نتایج حاصل از ارزیابی ریخت شناسی نشان داد، بیشترین اندازه طول برگ را اکوتیب های وری، ساری و کسوت نشان دادند و اکوتیب های ازنی و وری بیشترین اندازه طول ریشه را دارا بودند. همچنین، اکوتیپ های وری، کسوت، چهل چای و زنگلاب بالاترین میزان طول دم برگ و اکوتیپ چهل چای بالاترین تعداد برگ را نشان دادند. نتایج همبستگی صفات نشان داد که با افزایش ارتفاع از سطح دریا میزان طول برگ، عرض برگ، طول دم برگ، تعداد برگ، فلاونوئید برگ و فنل برگ گیاه بنفشه معطر کاهش می یابد. بهطورکلی اکوتیب های ساری، وری وکسوت در استان مازندران در اکثر ویژگی های عملکردی دارای ژنوتیب های برتر جهت کارهای اصلاحی در آینده می باشد.
In order to evaluate genetic diversity using phenotype and phytochemical properties of sweet violet (Viola odorata), 10 regions of Mazandaran and Golestan provinces were studied. Phytochemical traits such as phenol content, flavonoid, antioxidant capacity, chlorophyll a, chlorophyll b, total chlorophyll, and morphological characters of leaf size, petiole length, pedicel length, and root length were evaluated. Sari and Kosout genotypes had the highest chlorophyll b and total chlorophyll contents and the Sari ecotype had the highest flavonoid content. The highest level of antioxidant capacity was observed in Sari and Vari ecotypes. The ecotypes of Zanglab, Aq Cheshmeh, Tangeh Chehel Chai with lower height showed the highest total phenol contents. Results of morphological evaluation showed that ecotypes of Vari, Sar,i and kosout had the longest leaves and ecotypes of Azni and Vari had the longest roots. Also, ecotypes of Vari, kosout, Tangeh Chehel Chai, and Zanglab had the highest petiole length and Tangeh Chehel Chai ecotype had the highest leaf number. Correlation of traits showed that leaf height, leaf width, petiole length, leaf number, leaf flavonoid, and leaf phenol contents of sweet violet decreased with increasing altitude. In general, Sari, Vari, and Kosoot ecotypes from Mazandaran province had superior genotypes for most of the traits that could be used for future breeding programs.
Abbasnejad, R., Jabarzadeh, Z. and Razavi, M. (2017). The effect of different levels of light intensity on some physical and physiological characteristics of nightshade. Journal of Plant Research. 30 (2): 1-12.
Amin, G.h., Salehi, M.H., Yasa, N., Aynehchi, Y., Dehmoobed, A. and Emami, M. (2002). Screening of Iranian plants for antifungal activity: Part 2. Journal of Pharmaceutical Sciences. 10(2):78-89.
Arora, D.S. and Kaur, G.J. (2000). Antibacterial activity of some Indian medicinal plants. Journal of Natural Medicines, 61(3):313-17.
Arnon, D. (1947). Copper enzymes in isolated chloroplasts: Polyphenoloxidase in Beta vulgaris. Plant Physiology 24: 1-10.
Bernath, J. (2002). Strategies and recent achievements in selection of medicinal and aromatic plants. Proc. Int. Cont. on MAP. Acta Horticulture, 576: 65-68.
Cai, Y., Luo, Q., Sun, M. and Corke, H. (2004). Antioxidant activity and phenolic compounds of 112 traditional Chinese medicinal plant associated with anticancer'. Life Sciences. 74:2157–2184.
Carpenter Firoozjaei, M., Hemmati, K., Khorasani Nejad, S., Daraei Garmakhani, A. and Bagheri'fard, A. (2014). The effect of height on morphological and biochemical characteristics of nettle leaf (urtica dioica L.) in Mazandaran and Golestan provinces. Iranian Journal of Plant Ecophysiological Research. 9 (3): 1-11.
Chang, C., Yang, M., Wen, H. and Chern, J. (2002). Estimation of total flavonoid content in propolis by two complementary colorimetric methods. Journal of Food Drug Analysis, 10: 178-182.
Dou, H., Niu, G., Gu, M. and Masabni, Y. (2017). Effects of Light Quality on Growth and Phytonutrient Accumulation of Herbs under Controlled Environments. Horticulturae.36: 1-11.
Duke, J.A. (1985). Handbook of Medicinal Herbs. Boca Raton, FL: CRC Press;. 457.
Ebrahimzadeh, M.A., Nabavi, S.F., Nabavi, S.M. and Eslami, B. (2010). Antihemolytic and antioxidant activities of Allium paradoxum. Central European Journal of Biology. 5: 338-345.
Ghani, A., Nemati, S.H., Azizi, M., Saharkhiz, M.J. and Farsi, M. (2014). The study of extract biochemical variatons contents some of spearmint (Mentha spicata L.) Population. Journal of Horticulture Science. 4 (27): 433-443.
Hatamian, M., Arab, M. and Roozban, M.R. (2014). Photosynthetic and nonphotosynthetic pigments of two rose cultive under different light intensities. Agricultural Crop Management. 4 (16): 256-270.
Ireland, D.C., Colgrave, M.L. and Craik, D.J. (2006). A novel suite of cyclotides from Viola odorata: sequence variation and the implications for structure, function and stability. Biochemistry Journal, 400(1): 1-12.
Kaghazloo, Z. Hemti, Kh. And Khorasaninejad, S. (2017). The effect of altitude on some secondary metabolites of various organs of the sage plant (Sambucus ebulus L.) in three cities in Golestan province. Plant Environmental Physiology. (Iranian Plant Ecophysiology Research). 12 (3): 1-13.
Kohan'mu, M.A., Agha Alikhani, M., Rezazadeh, Sh. and Rejali, F. (2010). Investigation of growth potential and comparison of some agronomic and phytochemical characteristics of chamomile population of three natural habitats of Iran in Bushehr province. Iranian Journal of Horticulture Science and Technology, 11 (2): 98-78.
Karioti, A., Furlan, C., Vincieri, F.F. and Bilia, A.R. (2011). Analysis of the constituents and quality control of Viola odorata aqueous preparations by HPLC-DAD and HPLC-ESI-MS. Analytical and Bioanalytical Chemistry, 399(4): 1715-1723.
Kazemizadeh, Z., Habibi, Z. and Moradi, A. (2010). Investigation of the essential oil composition of Caapian Teucrium species (Teucrium hyrcanicum L.) in two different localities. Iranian Journal of Medicinal and Aromatic Plants Research, 9: 73-67.
Khanpour Ardestani, N., Sharifi, M. and Behmanesh, M. (2014). The effect of methyl jasmonate on the antioxidant activity of phenolic compounds and flavonoids in cell culture of Scrophularia straita Boiss. Journal of Plant Research. 27(5): 850-858.
Peng, L., Ru, M., Wang, B.Wang, Y., JingYu,, B. and Liang, Z. (2014). Genetic diversity assessment of a germplasm collection of Salvia miltiorrhiza Bunge. based on morphology, ISSR and SRAP markers. Biochemical Systematics and Ecology. 55: 84-92.
Lansari, A., Iezzoni, F. and Kester, D.E. (1994). Morphological variation within collections of Moroccan almond clones and Mediterranean and North American cultivars. Euphytica, 78: 27-41.
Lewis, W.H. and Elvin-Lewis, P.F. (1997). Medical Botany: Plants Affecting Man's Health. New Jersey, NJ: John Wiley & Sons. 389-90.
Mirzaei, A., Akbartabar, M., Sadeghi, H. and Sharifi, B. (2010). The Antioxidant Activities and total enolic of Artemisia Martima, Achillea Millefoliumand Matricaria Recutica. Iranian Society of Horticultural Science (IRSHS). 15(1): 243-252.
Mohammadnejad Ganji, S., M. Moradi, H., Ghanbari, A. and Akbarzadeh, M.A. (2017). Quantity and quality of secondary metabolite of lavender under the influence of ecological factor of altitude. New Findings in Life Sciences. 172: 4-.166.
Noormand Moaied, F., Bihamta, M.R., Tabaei-Aghdaei, S.R. and Naghavi, M.R. (2019). Study of morphological and phytochemical diversity among six (Satureja spp.) in East Azerbaijan province. Agricultural and Natural Resources Journal. 27(2): 252- 266.
Narimani, R., Moghaddam, M., Ghasemi Pir Balouti, A. and Shokouhi, D. (2017). Investigation of morphological diversity, phenol content and antioxidant activity of different populations of Nepeta nuda and Nepeta crassifolia in habitats of Ardabil and Azerbaijan provinces. Journal of Ecophytochemistry of Medicinal Plants. 5 (19): 22-13.
Nasibi, F., Kalantari, K. and Rashidi, M. (2003). Investigation of change in morphological and physiological parameter induced by UV-A, UV-B and UV-C of ultraviolet radiation in colza seedling (Brassica Napus). Research and Reconstruction. 16(3): 97-103.
Nowruz, A., Yousefzadeh, S., Asilan, K. and Mansoori'far, S. (2017). Evaluation of changes in essential oil, chlorophyll, carotenoids, anthocyanins and flavonoids of Mentha longifolia L. Hoda.subsp longifolia in different habitats of Marand. Journal of Ecophytochemical of Medicinal Plants. 5 (1): 64-52.
Omidbeigi, R. (2005). Production and processing of medicinal plant.- BahNashr. Mashhad. 1: 347
Pranting, M., Loov, C., Burman, R., Goransson, U. and Andersson, D.I. (2010). The cyclotide cycloviolacin O2 from Viola odorata has potent bactericidal activity against Gram-negative bacteria. Antimicrobial Agents and Chemotherapy; 65(9): 1964-71.
Pourmorad, F., Hosseinimehr, S.J. and Shahabimajd, N. (2006). Antioxidant activity, phenol and flavonoid contents of some selected iranian medicinal plants. African Journal of Biotechnology. 5 (11): 1142-1145.
Ramachandra, R.S. and Ravishankar, G.A. (2002). Plant cell cultures: Chemical factories of secondary metabolites. Biotechnology Advance. 20: 101-153.
Raeidehqi, H., Razmjoo, J., Sabzalian, M. and cheapness, A. (2015). The effect of shade on morpho-physiological characteristics and essential oil content of different genotypes of three species of mint. Plant Process and Function. 4 (13): 69-57.
Saiiad Aleian, M., Naderi, R., Fatahei Moghaddam, M.R. and Padasht Dehkaei, M.N. (2013). Evaluation of different plant populations Chelcheragh lily (Lilium ledebourii (Baker) Boiss.) using morphological characteristics and multivariate statistical methods. Iranian Journal of Horticultural Science. 44(4): 379-387.
Shokrollahi, S.h., Heshmati, Q.A. and Yousefzadeh, H. (2018). Investigation of phenol content, total flavonoids and antioxidant function of the rare plant Lilium ledebourii Boiss in northern Iran. Journal of Fasa University of Medical Sciences. 8 (2): 734-722.
Slinkard, K. and V.L. Singleton. (1977). Total phenol analysis:automation and comparison with manual methods. Amrican Journal of Enology and Viticulture, 28: 49-55.
Sonboli, A., Canaani, M., Yousefzadi, M. and Mojarrad, M. (2009). Comparisonof chemical composition and antibacterial effects of essential oil (Salvia hydrangea L.) in two different localities. Iranian Journal of Medicinal and Aromatic Plants Research, 9: 20-28.
Tatian, M.R. (2001). Sociological studies of plant (Phytosociology) summer pastures Hezarjarib-Behshahr.- Master s thesis. 128p.
Tawaha, K.H., Alali, F.Q., Gharaibeh, M., Mohammad, M. and Elimat, T.E. (2007). Antioxidant activity and total phenolic content of selected Jordanian plant species. Food Chemistry. 104: 1372– 1378.
Velioglu, Y.S., Mazza, G., Gao, L. and Oomah, B.D. (2000). Antioxidant activity and total phenolic in selected fruits, vegetables, and grain products. Agricultural and Food Chemistry. 46 (10): 4113-4127.
Zhang, J., Liao, B., Craik, D.J., Li, J.T.,
Hu, M. and Shu, W.S. (2009). Identification of two suites of cyclotide precursor genes from metallophyte Viola baoshanensis: cDNA sequence variation, alternative RNA splicing and potential cyclotide diversity. Gene. 431: 23–32.
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Abbasnejad, R., Jabarzadeh, Z. and Razavi, M. (2017). The effect of different levels of light intensity on some physical and physiological characteristics of nightshade. Journal of Plant Research. 30 (2): 1-12.
Amin, G.h., Salehi, M.H., Yasa, N., Aynehchi, Y., Dehmoobed, A. and Emami, M. (2002). Screening of Iranian plants for antifungal activity: Part 2. Journal of Pharmaceutical Sciences. 10(2):78-89.
Arora, D.S. and Kaur, G.J. (2000). Antibacterial activity of some Indian medicinal plants. Journal of Natural Medicines, 61(3):313-17.
Arnon, D. (1947). Copper enzymes in isolated chloroplasts: Polyphenoloxidase in Beta vulgaris. Plant Physiology 24: 1-10.
Bernath, J. (2002). Strategies and recent achievements in selection of medicinal and aromatic plants. Proc. Int. Cont. on MAP. Acta Horticulture, 576: 65-68.
Cai, Y., Luo, Q., Sun, M. and Corke, H. (2004). Antioxidant activity and phenolic compounds of 112 traditional Chinese medicinal plant associated with anticancer'. Life Sciences. 74:2157–2184.
Carpenter Firoozjaei, M., Hemmati, K., Khorasani Nejad, S., Daraei Garmakhani, A. and Bagheri'fard, A. (2014). The effect of height on morphological and biochemical characteristics of nettle leaf (urtica dioica L.) in Mazandaran and Golestan provinces. Iranian Journal of Plant Ecophysiological Research. 9 (3): 1-11.
Chang, C., Yang, M., Wen, H. and Chern, J. (2002). Estimation of total flavonoid content in propolis by two complementary colorimetric methods. Journal of Food Drug Analysis, 10: 178-182.
Dou, H., Niu, G., Gu, M. and Masabni, Y. (2017). Effects of Light Quality on Growth and Phytonutrient Accumulation of Herbs under Controlled Environments. Horticulturae.36: 1-11.
Duke, J.A. (1985). Handbook of Medicinal Herbs. Boca Raton, FL: CRC Press;. 457.
Ebrahimzadeh, M.A., Nabavi, S.F., Nabavi, S.M. and Eslami, B. (2010). Antihemolytic and antioxidant activities of Allium paradoxum. Central European Journal of Biology. 5: 338-345.
Ghani, A., Nemati, S.H., Azizi, M., Saharkhiz, M.J. and Farsi, M. (2014). The study of extract biochemical variatons contents some of spearmint (Mentha spicata L.) Population. Journal of Horticulture Science. 4 (27): 433-443.
Hatamian, M., Arab, M. and Roozban, M.R. (2014). Photosynthetic and nonphotosynthetic pigments of two rose cultive under different light intensities. Agricultural Crop Management. 4 (16): 256-270.
Ireland, D.C., Colgrave, M.L. and Craik, D.J. (2006). A novel suite of cyclotides from Viola odorata: sequence variation and the implications for structure, function and stability. Biochemistry Journal, 400(1): 1-12.
Kaghazloo, Z. Hemti, Kh. And Khorasaninejad, S. (2017). The effect of altitude on some secondary metabolites of various organs of the sage plant (Sambucus ebulus L.) in three cities in Golestan province. Plant Environmental Physiology. (Iranian Plant Ecophysiology Research). 12 (3): 1-13.
Kohan'mu, M.A., Agha Alikhani, M., Rezazadeh, Sh. and Rejali, F. (2010). Investigation of growth potential and comparison of some agronomic and phytochemical characteristics of chamomile population of three natural habitats of Iran in Bushehr province. Iranian Journal of Horticulture Science and Technology, 11 (2): 98-78.
Karioti, A., Furlan, C., Vincieri, F.F. and Bilia, A.R. (2011). Analysis of the constituents and quality control of Viola odorata aqueous preparations by HPLC-DAD and HPLC-ESI-MS. Analytical and Bioanalytical Chemistry, 399(4): 1715-1723.
Kazemizadeh, Z., Habibi, Z. and Moradi, A. (2010). Investigation of the essential oil composition of Caapian Teucrium species (Teucrium hyrcanicum L.) in two different localities. Iranian Journal of Medicinal and Aromatic Plants Research, 9: 73-67.
Khanpour Ardestani, N., Sharifi, M. and Behmanesh, M. (2014). The effect of methyl jasmonate on the antioxidant activity of phenolic compounds and flavonoids in cell culture of Scrophularia straita Boiss. Journal of Plant Research. 27(5): 850-858.
Peng, L., Ru, M., Wang, B.Wang, Y., JingYu,, B. and Liang, Z. (2014). Genetic diversity assessment of a germplasm collection of Salvia miltiorrhiza Bunge. based on morphology, ISSR and SRAP markers. Biochemical Systematics and Ecology. 55: 84-92.
Lansari, A., Iezzoni, F. and Kester, D.E. (1994). Morphological variation within collections of Moroccan almond clones and Mediterranean and North American cultivars. Euphytica, 78: 27-41.
Lewis, W.H. and Elvin-Lewis, P.F. (1997). Medical Botany: Plants Affecting Man's Health. New Jersey, NJ: John Wiley & Sons. 389-90.
Mirzaei, A., Akbartabar, M., Sadeghi, H. and Sharifi, B. (2010). The Antioxidant Activities and total enolic of Artemisia Martima, Achillea Millefoliumand Matricaria Recutica. Iranian Society of Horticultural Science (IRSHS). 15(1): 243-252.
Mohammadnejad Ganji, S., M. Moradi, H., Ghanbari, A. and Akbarzadeh, M.A. (2017). Quantity and quality of secondary metabolite of lavender under the influence of ecological factor of altitude. New Findings in Life Sciences. 172: 4-.166.
Noormand Moaied, F., Bihamta, M.R., Tabaei-Aghdaei, S.R. and Naghavi, M.R. (2019). Study of morphological and phytochemical diversity among six (Satureja spp.) in East Azerbaijan province. Agricultural and Natural Resources Journal. 27(2): 252- 266.
Narimani, R., Moghaddam, M., Ghasemi Pir Balouti, A. and Shokouhi, D. (2017). Investigation of morphological diversity, phenol content and antioxidant activity of different populations of Nepeta nuda and Nepeta crassifolia in habitats of Ardabil and Azerbaijan provinces. Journal of Ecophytochemistry of Medicinal Plants. 5 (19): 22-13.
Nasibi, F., Kalantari, K. and Rashidi, M. (2003). Investigation of change in morphological and physiological parameter induced by UV-A, UV-B and UV-C of ultraviolet radiation in colza seedling (Brassica Napus). Research and Reconstruction. 16(3): 97-103.
Nowruz, A., Yousefzadeh, S., Asilan, K. and Mansoori'far, S. (2017). Evaluation of changes in essential oil, chlorophyll, carotenoids, anthocyanins and flavonoids of Mentha longifolia L. Hoda.subsp longifolia in different habitats of Marand. Journal of Ecophytochemical of Medicinal Plants. 5 (1): 64-52.
Omidbeigi, R. (2005). Production and processing of medicinal plant.- BahNashr. Mashhad. 1: 347
Pranting, M., Loov, C., Burman, R., Goransson, U. and Andersson, D.I. (2010). The cyclotide cycloviolacin O2 from Viola odorata has potent bactericidal activity against Gram-negative bacteria. Antimicrobial Agents and Chemotherapy; 65(9): 1964-71.
Pourmorad, F., Hosseinimehr, S.J. and Shahabimajd, N. (2006). Antioxidant activity, phenol and flavonoid contents of some selected iranian medicinal plants. African Journal of Biotechnology. 5 (11): 1142-1145.
Ramachandra, R.S. and Ravishankar, G.A. (2002). Plant cell cultures: Chemical factories of secondary metabolites. Biotechnology Advance. 20: 101-153.
Raeidehqi, H., Razmjoo, J., Sabzalian, M. and cheapness, A. (2015). The effect of shade on morpho-physiological characteristics and essential oil content of different genotypes of three species of mint. Plant Process and Function. 4 (13): 69-57.
Saiiad Aleian, M., Naderi, R., Fatahei Moghaddam, M.R. and Padasht Dehkaei, M.N. (2013). Evaluation of different plant populations Chelcheragh lily (Lilium ledebourii (Baker) Boiss.) using morphological characteristics and multivariate statistical methods. Iranian Journal of Horticultural Science. 44(4): 379-387.
Shokrollahi, S.h., Heshmati, Q.A. and Yousefzadeh, H. (2018). Investigation of phenol content, total flavonoids and antioxidant function of the rare plant Lilium ledebourii Boiss in northern Iran. Journal of Fasa University of Medical Sciences. 8 (2): 734-722.
Slinkard, K. and V.L. Singleton. (1977). Total phenol analysis:automation and comparison with manual methods. Amrican Journal of Enology and Viticulture, 28: 49-55.
Sonboli, A., Canaani, M., Yousefzadi, M. and Mojarrad, M. (2009). Comparisonof chemical composition and antibacterial effects of essential oil (Salvia hydrangea L.) in two different localities. Iranian Journal of Medicinal and Aromatic Plants Research, 9: 20-28.
Tatian, M.R. (2001). Sociological studies of plant (Phytosociology) summer pastures Hezarjarib-Behshahr.- Master s thesis. 128p.
Tawaha, K.H., Alali, F.Q., Gharaibeh, M., Mohammad, M. and Elimat, T.E. (2007). Antioxidant activity and total phenolic content of selected Jordanian plant species. Food Chemistry. 104: 1372– 1378.
Velioglu, Y.S., Mazza, G., Gao, L. and Oomah, B.D. (2000). Antioxidant activity and total phenolic in selected fruits, vegetables, and grain products. Agricultural and Food Chemistry. 46 (10): 4113-4127.
Zhang, J., Liao, B., Craik, D.J., Li, J.T.,
Hu, M. and Shu, W.S. (2009). Identification of two suites of cyclotide precursor genes from metallophyte Viola baoshanensis: cDNA sequence variation, alternative RNA splicing and potential cyclotide diversity. Gene. 431: 23–32.