کاربرد هورمون بتااسترادیول با هدف افزایش تحمل تنش به شوری در ژنوتیپ های سیبزمینی (Solanum tuberosum L)
محورهای موضوعی : اکوفیزیولوژی گیاهان زراعیفهیمه جی دار 1 , رسول اصغری زکریا 2 , ناصر زارع 3 , داوود حسن پناه 4 , لیلا غفارزاده نمازی 5
1 - دانشجوی دکتری، گروه تولید و ژنتیک گیاهی، دانشگاه محقق اردبیلی، اردبیل، ایران
2 - استاد گروه تولید و ژنتیک گیاهی، دانشگاه محقق اردبیلی، اردبیل، ایران
3 - استاد گروه تولید و ژنتیک گیاهی، دانشگاه محقق اردبیلی، اردبیل، ایران
4 - عضو هیات علمی بخش تحقیقات زراعی و باغی، مرکز تحقیقات و آموزش کشاورزی و منابع طبیعی استان اردبیل (مغان)، سازمان تحقیقات، آموزش و ترویج کشاورزی، اردبیل، ایران
5 - استادیار، گروه علوم گیاهی و گیاهان دارویی، دانشگاه محقق اردبیلی، دانشکده کشاورزی مشکین شهر، مشکینشهر، ایران
کلید واژه: نشاسته, آنزیمهای آنتیاکسیدان, هورمونهای استروئیدی, غدهچه,
چکیده مقاله :
این آزمایش به منظور بررسی تأثیر کاربرد هورمون بتااسترادیول بر افزایش تحمل به تنش شوری ژنوتیپهای مختلف سیبزمینی، به صورت فاکتوریل اسپلیت پلات بر پایه طرح بلوکهای کامل تصادفی با سه تکرار در دانشگاه محقق اردبیلی در سال 1398 انجام شد. تیمارهای تنش شوری شامل سه سطح (صفر، 50 و 100 میلیمولار کلرید سدیم) و هورمون بتااسترادیول در سه سطح (صفر، 12-10 و 6-10 مولار) به صورت فاکتوریل در کرتهای اصلی، و 10 ژنوتیپ سیبزمینی در کرت های فرعی بودند. نتایج نشان داد ارتفاع بوته، تعداد و وزن غده چه در بوته، وزن متوسط غدهچه، محتوی آنزیمهای آنتیاکسیدان در ژنوتیپهای مورد بررسی تحت تاثیر کاربرد هورمون بتااسترادیول قرار گرفتند. به طوری که با افزایش میزان مصرف بتااسترادیول از 12-10 به 6-10 مولار، تعداد و وزن غدهچه در بیشتر ژنوتیپ های مورد مطالعه افزایش یافت ولی مقدار این افزایش در بین ژنوتیپها متفاوت بود. ژنوتیپ های G5 و G6 به ترتیب با میانگین 7.85 و 7.83غده چه، بالاترین تعداد غدهچه در بوته را در سطح 6-10 مولار بتااسترادیول به خود اختصاص دادند. کمترین مقدار صفت مذکور نیز با میانگین 3.66 غده چه بدون اختلاف معنیدار با ژنوتیپهای G8 و G9 به ژنوتیپ G10 تعلق داشت. با افزایش سطح شوری میزان آنزیمهای سوپراکسید دیسموتاز، کاتالاز و پلی فنل اکسیداز و قندهای محلول افزایش یافت. در هر سه سطح شوری مصرف بتااسترادیول به طور معنیداری باعث افزایش میزان این آنزیمها شد. بالاترین میزان این آنزیمها در سطح شوری 100 میلی مولار و با مصرف 12-10 یا 6-10 مولار بتااسترادیول مشاهده شد. در این تحقیق کاربرد بتااسترادیول بسته به ژنوتیپ توانست اثر شوری را بر خصوصیات کمّی و کیفی سیبزمینی تعدیل نماید. در کل، در این مطالعه ژنوتیپهای G5 و G6 در شرایط تنش شوری به طور نسبی از تعداد و وزن غدهچه در بوته بالایی برخوردار بودند، بنابراین گزینش این ژنوتیپها برای برنامههای به نژادی آتی توصیه می شود. همچنین، این دو ژنوتیپ در سطوح 12-10 و 6-10 مولار بتااسترادیول بیشترین تعداد و وزن غده چه در بوته را به خود اختصاص دادند که نشان میدهد این ژنوتیپها در مقایسه با دیگر ژنوتیپهای مورد بررسی از پتانسیل ژنتیکی بالایی برای مصرف این هورمون برخوردارند.
This experiment was performed to investigate the effect of beta estradiol hormone application on increasing salinity stress tolerance of different potato genotypes in a factorial split plot experiment based on a randomized complete block design with three replications in Mohaghegh Ardabili University in 2020. Salinity stresses with three levels (0, 50 and 100 mM sodium chloride) and beta-estradiol also with three levels (0, 10-12 and 6-10 M) were assigned to main plots, and 10 potato genotypes to subplots. The results revealed that plant height, number and weight of minituber per plant, average tuber weight and content of antioxidant enzymes in the studied genotypes showed a positive response to beta-estradiol. Thus, with increasing the amount of beta-estradiol application from 10-12 to 10-6 M, the number and weight of minitubers increased in most of the studied genotypes, but the amount of this increase was varied between genotypes. G5 and G6 genotypes with an average of 7.85 and 7.83 minitubers had the highest number of tubers per plant at 10-6 M beta-estradiol, respectively. The lowest value of this trait belonged to G10 genotype with an average of 3.66 minitubers, without significant differences with those of G8 and G9 genotypes. With increasing salinity level, the enzymes of superoxide dismutase, catalase and polyphenol oxidase and also soluble sugars were increased. In all of three salinity levels, beta-estradiol application significantly increased the levels of these enzymes. The highest levels of these enzymes were observed at salinity level of 100 mM with the use of 10-12 or 6-10 M beta estradiol. In this study, the use of beta-estradiol, depending on the genotype, was able to moderate the effect of salinity on the quantitative and qualitative characteristics of potatoe tubers. Overall, in this study, G5 and G6 genotypes had relatively high minituber number and weight per plant under salinity stress conditions, Thus, selection of these genotypes is recommended for future breeding programs. Also, these two genotypes had the highest number and weight of minitubers per plant at 10-12 and 10-6 M beta estradiol, which indicates that these genotypes have a high genetic potential for consumption of this hormone as compared to other genotypes.
Aghaei, K., A.A. Ehsanpour, and S. Komatsu. 2009. Potato responds to salt stress by increased activity of antioxidant enzymes. Journal of Integrative Plant Biology. 51(12): 1095–1103.
Ahmed, A.A., N. Koçak Şahin, G. Akdoğan, C. Yaman, D. Köm, and S. Uranbey. 2020. Variability in salinity stress tolerance of potato (Solanum tuberosum L.) varieties using in vitro screening. Ciência e Agrotecnologia. 44: 1-14.
Amerian, M., and M. Esna-Ashari. 2017. Effect of different levels of salinity on some physiological and cells-growth characteristics in three potato (Solanum tubrosum) cultivars in vitro. Plant Production Technology. 17(1): 209-225. (In Persian).
Asada, K., M. Takahashi, and M. Nagate. 1974. Assay and inhibitors of spinach superoxide dismutase. Agricultural and Biological Chemistry. 38: 471-473.
Bates, L.S., R.P. Waldren, and I.D. Teare. 1973. Rapid determination of free proline for water stress Plant and Soil. 39: 205-207.
Bradford, M.M. 1976. A rapid and sensitive method for the qualify cation of microgram quantities of protein utilizing the principle of protein dye binding. Analytical Biochemistry. 72: 248-254.
Brown, C. 2006. The effects of estrogen on the growth and tuberization of potato plants (Solanum tuberosum ‘Iwa’) grown in liquid tissue culture media. A Thesis for the Degree of Master of Science in Plant Biotechnology. University of Canterbury School of Biological Sciences. 128P
Bruce, W., O. Folkerts, C. Garnaat, O. Crasta, B. Roth, and B. Bowen. 2000. Expression profiling of the maize flavonoid pathway genes controlled by estradiol-inducible transcription factors CRC and P. Plant Cell. 12: 65–80.
Bündig, C., H. Vu, P. Meise, S. Seddig, and A. Schum. 2016. Winkelmann variability in osmotic stress tolerance of starch potato genotypes (Solanum tuberosum L.) as revealed by an in vitro screening: Role of proline, osmotic adjustment and drought response in pot trials. Journal of Agronomy and Crop Science. 203: 206-218.
Carreau, S., C. Delalande, D. Silandre, Bourguiba, and S. Lambard. 2004. Aromatase and estrogen receptors in male reproduction. Molecular and Cellular Endocrinology. 246(1): 65-68.
Chaoui, A., and E. El Ferjani. 2013. β-Estradiol protects embryo growth from heavy-metal toxicity in germinating lentil seeds. Journal of Plant Growth Regulation. 32 (1): 1-16.
Charfeddine, M., Charfeddine, I. Ghazala, D. Bouaziz, and A. Radhia Gargouri Bouzid. 2019. Investigation of the response to salinity of transgenic potato plants overexpressing the transcription factor StERF94. Journal of Biosciences. 44(141): 2-16.
Chinsamy, M., M.G. Kulkarni, and S. Van. 2013. Gardenwaste- vermicompost leachate alleviates salinity stress in tomato seedlings by mobilizing salt tolerance mechanisms. Plant Growth Regulation. 71: 41–47.
Çulha, Ş., and H. Çakirlar, 2011. The effect of salinity on plants and salt tolerance mechanisms. Afyon Kocatepe Üniversitesi Fen Bilimleri Dergisi. 11: 11-34.
Dumlupinar, R., M. Genisel Erdal, T. Korkut, M.S. Taspinar, and T. Taskin. 2011. Effects of progesterone, β-estradiol and androsterone on the changes of inorganic element content in barley leaves. Biological Trace Element Research. 143: 1740–1745.
Eisenbarth, D.A., and A.R. Weig. 2005. Dynamics of aquaporins and water relations during hypocotyl elongation in Ricinus communis seedlings. Journal of Experimental Botany. 56: 1831–1842.
Erdal, S. 2011. Alleviation of salt stress in wheat seedlings by mammalian sex hormones. Journal of the Science of Food and Agriculture. 92: 1411-1416.
Erdal, S. 2012a. Exogenous mammalian sex hormones mitigate inhibition in growth by enhancing antioxidant activity and synthesis reactions in germinating maize seeds under salt stress. Science of Food and Agriculture. 92: 839–843.
Erdal, S. 2012b. Alleviation of salt stress in wheat seedlings by mammalian sex hormones. Science of Food and Agriculture. 92: 1411–1416.
Erdal, S., and R. Dumlupinar. 2011. Mammalian sex hormones stimulate antioxidant system and enhance growth of chickpea plants. Acta Physiologiae Plantarum. 33: 1011–1017.
Faried, H.N., C. Muhammad Ayyub, M. Amjad Rashid Ahmed, F. MasoudWattoo, and M. Butt. 2017. Salicylic acid confers salt tolerance in potato plants by improving water relations, gaseous exchange, antioxidant activities and osmoregulation. Science of Food and Agriculture. 97 (6): 1868-1875.
Genisel, M., H. Turk, and S. Erdal. 2013. Exogenous progesterone application protects chickpea seedlings against chilling-induced oxidative stress. Acta Physiologiae Plantarum. 35(1): 241–251.
Gu, F., R. Hata, K. Toku, L.Yang, Y.J. Ma, and N. Maeda. 2003. Testosterone up-regulates aquaporin-4 expression in cultured astrocytes. Journal of Neuroscience Research. 72: 709-715.
Hassanpanah, D., and R. Asghari Zakaria. 2018. Evaluation of radiated potato genotypes with gamma rays in water deficit stress. Agricultural Science and Sustainable Production. 28(2): 107-122. (In Persian).
Homayoun, H., P. Mehrabi, and M. Sam Daliri. 2011. Study of salinity stress effect on two commercial varieties of potato (Solanum tuberosum) after transmitting to green house from in vitro culture. American-Eurasian Journal of Agricultural & Environmental Sciences. 11(5): 725-728.
Isayenkov, S.V., and F.J.M. Maathuis. 2019. Plant salinity stress: Many unanswered questions remain. Frontiers in Plant Science. 10(80): 1-11.
Janeczko, A. 2000. Influence of selected steroids on plant physiological processes especially flowering induction. Ph.D. Thesis, Agriculture University, Krakow.
Janeczko, A., and A. Skoczowski. 2005. Mammalian sex hormones in plants. Folia Histochemica ET Cytobiologica. 43(2): 71-79.
Janeczko, A., W. Filek, J. Biesaga-Kościelniak, I. Marcińska, and Z. Janeczko. 2003. The influence of animal sex hormones on the induction of flowering in Arabidopsis thaliana: comparison with the effect of 24-Epibrassinolide. Plant Cell, Tissue and Organ Culture. 72: 147-151
Jensen, C.R., A. Battilani, F. Plauborg, G. Psarras, K. Chartzoulakis, F. Janowiak, R. Stikic, Z. Jovanovic, G. Li, X. Qi, F. Liu, S.E. Jacobsen, and M.N. Andersen. 2010. Deficit irrigation based on drought tolerance and root signalling in potatoes and tomatoes. Agricultural Water Management. 98(3): 403-413.
Jha, G., P. Choudhary, and R. Sharda Manuel Tejada Moral. 2017. Comparative effects of saline water on yield and quality of potato under drip and furrow irrigation. Cogent Food and Agriculture. 3(1): 2-14.
Jing, X., Hou, Y. Lu, S. Deng, N. Li, R. Zhao, J. Sun, Y. Wang, Y. Han, T. Lang, M. Ding, X. Shen, and S. Chen. 2015. Overexpression of copper/zinc superoxide dismutase from mangrove Kandelia candel in tobacco enhances salinity tolerance by the reduction of reactive oxygen species in chloroplast. Frontiers in Plant Science. 6(23): 1-13.
Kafi, M., J. Nabati, B. Saadatian, A. Oskoiyan, and E. Boroumand Rezazadeh. 2016. Adjustment of salinity effect by silicon compounds (nano and micro) and potassium on in potato. 15th National Iranian Crop Science Congress. Sep. 4-6, 2018. Karaj, Iran. (In Persian).
Kar, M., and D. Mishra .1976. Catalase, peroxidase, and polyphenoloxidase activities during rice leaf senescence. Plant Physiology. 57(2): 315-319.
Khalid, A., and F. Aftab. 2016. Effect of exogenous application of 24-epibrassinolideon growth, protein contents, and antioxidant enzyme activities of in vitro-grown (Solanum tuberosum) under salt stress. In Vitro Cellular and Developmental Biology-Plant. 52: 81-91.
Krauss, S., W.H. Schnitzler, J. Grassmann, and M. Woitke. The influence of different electrical conductivity values in a simplified recirculating soilless system on inner and outer fruit quality characteristics of tomato. Journal of Agricultural and Food Chemistry. 54: 441–448.
Levy, D., and G.C.C. Tai. 2013. Differential response of potatoes (Solanum tuberosum) to salinity in an arid environment and field performance of the seed tubers grown with fresh water in the following season. Agricultural Water Management. 116: 122–127.
McCready, R.M., J. Guggolz, V. Silviera, and H.S. Owens. 1950. Determination of starch and amylose in vegetables. Analytical Chemistry. 22: 1156–1158.
Moghaddaszadeh-Ahrabi, M., R. Asghari Zakaria, D. Hassanpanah, and Zare. 2018. Evaluation of agronomic traits and yield stability in several potato (Solanum tuberosum L.) genotypes. Journal of Crop Ecophysiology.12(45): 153-170 (In Persian).
Mohammadi, Z., A. Motallebi Azar, F. Zaare-Nahandi, A. Tarinejad, and G. Gohari. Effect of sodium nitroprusside on growth, physiological and biochemical characters of Solanum tuberosum cv. Agria under salinity stress on in vitro condition. Journal of Plant Production Research. 26(1): 155-167. (In Persian).
Morillon, R., M. Catterou, R.S. Sangwan, B.S. Sangwan, and J.P. Lassalles. 2001. Brassinolide may control aquaporin activities in Arabidopsis thaliana. Planta. 212: 199-204.
Morsy, M.R., L. Jouve, J.F. Hausman, L. Hoffmann, and J.M. Stewart. 2007. Alteration of oxidative and carbohydrate metabolism under abiotic stress in two rice (Oryza sativa) genotypes contrasting in chilling tolerance. Journal of Plant Physiology. 164: 157–67.
Munira, S., M.M. Hossain, M. Zakaria, J.U. Ahmed, and M.M. Islam. 2015. Evaluation of potato varieties against salinity stress in Bangladesh. International Journal of Plant & Soil Science. 6(2): 73-81.
Nouri, A., A. Ahmad Nezami, M. Kafi, and D. Hassanpanah. 2016. Evaluation of water deficit tolerance of 10 potato cultivars based on some physiological traits and (Solanum tuberosum) tuber yield in Ardabil region. Journal of Crop Ecophysiology. 1(37): 243-268. (In Persian).
Nozari, E., R. Asghari-Zakaria, S. Jahanbakhsh, and N. Zare. 2018. The effect of steroidal testosterone hormone on seedling growth, antioxidant enzymes activity and callus induction in German chamomile (Matricaria chamomilla). Journal of Crop Breeding. 10(27): 31-38. (In Persian).
Parihar, P., Singh, R. Singh, V. Pratap Singh, and S. Mohan Prasad. 2015. Effect of salinity stress on plants and its tolerance strategies: A review. Environmental Science and Pollution Research. 22(6): 4056-4075.
Patade, V.Y., P. Suprasanna, and V.A. Bapat. 2006. Selection for abiotic (salinity and drought) stress tolerance and molecular characterization of tolerance lines in sugarcane. The National Conference on Biotechnological Aspects towards Cultivation, Utilization and Disease Management of Plants. Issue. 273.
Perez-Gomez, J.J., M. Abud-Archila, J. Jose Villalobos-Maldonado, S. Enciso-Saenz, H. Hernandez deLeon, V.M. Ruiz-Valdiviezo, and F.A. Gutierrez-Miceli. 2017. Vermicompost and vermiwash minimized the influence of salinity stress on growth parameters in potato plants. Compost Science & Utilization. 12: 1-7.
Rykaczewska, K. 2013. The impact of high temperature during growing season on potato cultivars with different response to environmental stresses. American Journal of Plant Sciences. 4: 2386–2393.
Schlegel, H.G. 1956. Die verwertung organischer sauren durch chlorella in lincht. Planta. 47: 510-515.
Shibli, R.A., M. Kushad, G.G. Yousef, and M. Lila, 2007. Physiological and biochemical responses of tomato micro shoots to induced salinity stress with associated ethylene accumulation. Plant Growth Regulation. 51: 159-169.
Shimazaki, T., T. Endo, M. Kasuga, K. Yamaguchi-Shinozaki, K.N. Watanabe, and A. Kikuchi. 2017. Evaluation of the yield of abiotic-stress-tolerant tDREB1A transgenic potato under saline conditions in advance of field trials. Breeding Science. 66: 703–710.
Simersky, R., O. Nova´, K.D.A. Morris, V. Pouzar, and M. Strnad .2009. Identification and quantification of several mammalian steroid hormones in plants by UPLC–MS/MS. Journal of Plant Growth Regulation. 28: 125–136.
Sudhersan, C., Jibi Manuel, J. Ashkanani, and A. Al-Ajeel. 2012. In vitro screening of potato cultivars for salinity tolerance. American-Eurasian Journal of Sustainable Agriculture. 6(4): 344-348.
Uranbey, S., D. Köm, Akdoğan, H.A.A. Ahmed, N. Koçak, and M.E. Kara. 2017. The effects of salt stress on expression of Asg1 gene related stomatal resistance in potato. Mediterranean Agricultural Sciences. 30(3): 235-238.
Velikova, V., I. Yordanov, and A. Edreva .2000. Oxidative stress and some antioxidant systems in acid rain treated bean plants. Protective role of exogenous polyamines. Plant Science. 151: 59-66.
Wang, L., Y. Liu, D. Li, S. Feng, J. Yang, J. Zhang, J. Zhang, Wang, and Y. Gan. 2017. Improving salt tolerance in potato through overexpression of AtHKT1 gene. BMC Plant Biology. 9(357): 1-57.
Zaman, S.M., M. Ali, A. Muhammad, and I. Hussain. 2015. In vitro screening of salt tolerance in potato (Solanum tuberosum L.) varieties. Sarhad Journal of Agriculture. 31(2): 106-113.
_||_Aghaei, K., A.A. Ehsanpour, and S. Komatsu. 2009. Potato responds to salt stress by increased activity of antioxidant enzymes. Journal of Integrative Plant Biology. 51(12): 1095–1103.
Ahmed, A.A., N. Koçak Şahin, G. Akdoğan, C. Yaman, D. Köm, and S. Uranbey. 2020. Variability in salinity stress tolerance of potato (Solanum tuberosum L.) varieties using in vitro screening. Ciência e Agrotecnologia. 44: 1-14.
Amerian, M., and M. Esna-Ashari. 2017. Effect of different levels of salinity on some physiological and cells-growth characteristics in three potato (Solanum tubrosum) cultivars in vitro. Plant Production Technology. 17(1): 209-225. (In Persian).
Asada, K., M. Takahashi, and M. Nagate. 1974. Assay and inhibitors of spinach superoxide dismutase. Agricultural and Biological Chemistry. 38: 471-473.
Bates, L.S., R.P. Waldren, and I.D. Teare. 1973. Rapid determination of free proline for water stress Plant and Soil. 39: 205-207.
Bradford, M.M. 1976. A rapid and sensitive method for the qualify cation of microgram quantities of protein utilizing the principle of protein dye binding. Analytical Biochemistry. 72: 248-254.
Brown, C. 2006. The effects of estrogen on the growth and tuberization of potato plants (Solanum tuberosum ‘Iwa’) grown in liquid tissue culture media. A Thesis for the Degree of Master of Science in Plant Biotechnology. University of Canterbury School of Biological Sciences. 128P
Bruce, W., O. Folkerts, C. Garnaat, O. Crasta, B. Roth, and B. Bowen. 2000. Expression profiling of the maize flavonoid pathway genes controlled by estradiol-inducible transcription factors CRC and P. Plant Cell. 12: 65–80.
Bündig, C., H. Vu, P. Meise, S. Seddig, and A. Schum. 2016. Winkelmann variability in osmotic stress tolerance of starch potato genotypes (Solanum tuberosum L.) as revealed by an in vitro screening: Role of proline, osmotic adjustment and drought response in pot trials. Journal of Agronomy and Crop Science. 203: 206-218.
Carreau, S., C. Delalande, D. Silandre, Bourguiba, and S. Lambard. 2004. Aromatase and estrogen receptors in male reproduction. Molecular and Cellular Endocrinology. 246(1): 65-68.
Chaoui, A., and E. El Ferjani. 2013. β-Estradiol protects embryo growth from heavy-metal toxicity in germinating lentil seeds. Journal of Plant Growth Regulation. 32 (1): 1-16.
Charfeddine, M., Charfeddine, I. Ghazala, D. Bouaziz, and A. Radhia Gargouri Bouzid. 2019. Investigation of the response to salinity of transgenic potato plants overexpressing the transcription factor StERF94. Journal of Biosciences. 44(141): 2-16.
Chinsamy, M., M.G. Kulkarni, and S. Van. 2013. Gardenwaste- vermicompost leachate alleviates salinity stress in tomato seedlings by mobilizing salt tolerance mechanisms. Plant Growth Regulation. 71: 41–47.
Çulha, Ş., and H. Çakirlar, 2011. The effect of salinity on plants and salt tolerance mechanisms. Afyon Kocatepe Üniversitesi Fen Bilimleri Dergisi. 11: 11-34.
Dumlupinar, R., M. Genisel Erdal, T. Korkut, M.S. Taspinar, and T. Taskin. 2011. Effects of progesterone, β-estradiol and androsterone on the changes of inorganic element content in barley leaves. Biological Trace Element Research. 143: 1740–1745.
Eisenbarth, D.A., and A.R. Weig. 2005. Dynamics of aquaporins and water relations during hypocotyl elongation in Ricinus communis seedlings. Journal of Experimental Botany. 56: 1831–1842.
Erdal, S. 2011. Alleviation of salt stress in wheat seedlings by mammalian sex hormones. Journal of the Science of Food and Agriculture. 92: 1411-1416.
Erdal, S. 2012a. Exogenous mammalian sex hormones mitigate inhibition in growth by enhancing antioxidant activity and synthesis reactions in germinating maize seeds under salt stress. Science of Food and Agriculture. 92: 839–843.
Erdal, S. 2012b. Alleviation of salt stress in wheat seedlings by mammalian sex hormones. Science of Food and Agriculture. 92: 1411–1416.
Erdal, S., and R. Dumlupinar. 2011. Mammalian sex hormones stimulate antioxidant system and enhance growth of chickpea plants. Acta Physiologiae Plantarum. 33: 1011–1017.
Faried, H.N., C. Muhammad Ayyub, M. Amjad Rashid Ahmed, F. MasoudWattoo, and M. Butt. 2017. Salicylic acid confers salt tolerance in potato plants by improving water relations, gaseous exchange, antioxidant activities and osmoregulation. Science of Food and Agriculture. 97 (6): 1868-1875.
Genisel, M., H. Turk, and S. Erdal. 2013. Exogenous progesterone application protects chickpea seedlings against chilling-induced oxidative stress. Acta Physiologiae Plantarum. 35(1): 241–251.
Gu, F., R. Hata, K. Toku, L.Yang, Y.J. Ma, and N. Maeda. 2003. Testosterone up-regulates aquaporin-4 expression in cultured astrocytes. Journal of Neuroscience Research. 72: 709-715.
Hassanpanah, D., and R. Asghari Zakaria. 2018. Evaluation of radiated potato genotypes with gamma rays in water deficit stress. Agricultural Science and Sustainable Production. 28(2): 107-122. (In Persian).
Homayoun, H., P. Mehrabi, and M. Sam Daliri. 2011. Study of salinity stress effect on two commercial varieties of potato (Solanum tuberosum) after transmitting to green house from in vitro culture. American-Eurasian Journal of Agricultural & Environmental Sciences. 11(5): 725-728.
Isayenkov, S.V., and F.J.M. Maathuis. 2019. Plant salinity stress: Many unanswered questions remain. Frontiers in Plant Science. 10(80): 1-11.
Janeczko, A. 2000. Influence of selected steroids on plant physiological processes especially flowering induction. Ph.D. Thesis, Agriculture University, Krakow.
Janeczko, A., and A. Skoczowski. 2005. Mammalian sex hormones in plants. Folia Histochemica ET Cytobiologica. 43(2): 71-79.
Janeczko, A., W. Filek, J. Biesaga-Kościelniak, I. Marcińska, and Z. Janeczko. 2003. The influence of animal sex hormones on the induction of flowering in Arabidopsis thaliana: comparison with the effect of 24-Epibrassinolide. Plant Cell, Tissue and Organ Culture. 72: 147-151
Jensen, C.R., A. Battilani, F. Plauborg, G. Psarras, K. Chartzoulakis, F. Janowiak, R. Stikic, Z. Jovanovic, G. Li, X. Qi, F. Liu, S.E. Jacobsen, and M.N. Andersen. 2010. Deficit irrigation based on drought tolerance and root signalling in potatoes and tomatoes. Agricultural Water Management. 98(3): 403-413.
Jha, G., P. Choudhary, and R. Sharda Manuel Tejada Moral. 2017. Comparative effects of saline water on yield and quality of potato under drip and furrow irrigation. Cogent Food and Agriculture. 3(1): 2-14.
Jing, X., Hou, Y. Lu, S. Deng, N. Li, R. Zhao, J. Sun, Y. Wang, Y. Han, T. Lang, M. Ding, X. Shen, and S. Chen. 2015. Overexpression of copper/zinc superoxide dismutase from mangrove Kandelia candel in tobacco enhances salinity tolerance by the reduction of reactive oxygen species in chloroplast. Frontiers in Plant Science. 6(23): 1-13.
Kafi, M., J. Nabati, B. Saadatian, A. Oskoiyan, and E. Boroumand Rezazadeh. 2016. Adjustment of salinity effect by silicon compounds (nano and micro) and potassium on in potato. 15th National Iranian Crop Science Congress. Sep. 4-6, 2018. Karaj, Iran. (In Persian).
Kar, M., and D. Mishra .1976. Catalase, peroxidase, and polyphenoloxidase activities during rice leaf senescence. Plant Physiology. 57(2): 315-319.
Khalid, A., and F. Aftab. 2016. Effect of exogenous application of 24-epibrassinolideon growth, protein contents, and antioxidant enzyme activities of in vitro-grown (Solanum tuberosum) under salt stress. In Vitro Cellular and Developmental Biology-Plant. 52: 81-91.
Krauss, S., W.H. Schnitzler, J. Grassmann, and M. Woitke. The influence of different electrical conductivity values in a simplified recirculating soilless system on inner and outer fruit quality characteristics of tomato. Journal of Agricultural and Food Chemistry. 54: 441–448.
Levy, D., and G.C.C. Tai. 2013. Differential response of potatoes (Solanum tuberosum) to salinity in an arid environment and field performance of the seed tubers grown with fresh water in the following season. Agricultural Water Management. 116: 122–127.
McCready, R.M., J. Guggolz, V. Silviera, and H.S. Owens. 1950. Determination of starch and amylose in vegetables. Analytical Chemistry. 22: 1156–1158.
Moghaddaszadeh-Ahrabi, M., R. Asghari Zakaria, D. Hassanpanah, and Zare. 2018. Evaluation of agronomic traits and yield stability in several potato (Solanum tuberosum L.) genotypes. Journal of Crop Ecophysiology.12(45): 153-170 (In Persian).
Mohammadi, Z., A. Motallebi Azar, F. Zaare-Nahandi, A. Tarinejad, and G. Gohari. Effect of sodium nitroprusside on growth, physiological and biochemical characters of Solanum tuberosum cv. Agria under salinity stress on in vitro condition. Journal of Plant Production Research. 26(1): 155-167. (In Persian).
Morillon, R., M. Catterou, R.S. Sangwan, B.S. Sangwan, and J.P. Lassalles. 2001. Brassinolide may control aquaporin activities in Arabidopsis thaliana. Planta. 212: 199-204.
Morsy, M.R., L. Jouve, J.F. Hausman, L. Hoffmann, and J.M. Stewart. 2007. Alteration of oxidative and carbohydrate metabolism under abiotic stress in two rice (Oryza sativa) genotypes contrasting in chilling tolerance. Journal of Plant Physiology. 164: 157–67.
Munira, S., M.M. Hossain, M. Zakaria, J.U. Ahmed, and M.M. Islam. 2015. Evaluation of potato varieties against salinity stress in Bangladesh. International Journal of Plant & Soil Science. 6(2): 73-81.
Nouri, A., A. Ahmad Nezami, M. Kafi, and D. Hassanpanah. 2016. Evaluation of water deficit tolerance of 10 potato cultivars based on some physiological traits and (Solanum tuberosum) tuber yield in Ardabil region. Journal of Crop Ecophysiology. 1(37): 243-268. (In Persian).
Nozari, E., R. Asghari-Zakaria, S. Jahanbakhsh, and N. Zare. 2018. The effect of steroidal testosterone hormone on seedling growth, antioxidant enzymes activity and callus induction in German chamomile (Matricaria chamomilla). Journal of Crop Breeding. 10(27): 31-38. (In Persian).
Parihar, P., Singh, R. Singh, V. Pratap Singh, and S. Mohan Prasad. 2015. Effect of salinity stress on plants and its tolerance strategies: A review. Environmental Science and Pollution Research. 22(6): 4056-4075.
Patade, V.Y., P. Suprasanna, and V.A. Bapat. 2006. Selection for abiotic (salinity and drought) stress tolerance and molecular characterization of tolerance lines in sugarcane. The National Conference on Biotechnological Aspects towards Cultivation, Utilization and Disease Management of Plants. Issue. 273.
Perez-Gomez, J.J., M. Abud-Archila, J. Jose Villalobos-Maldonado, S. Enciso-Saenz, H. Hernandez deLeon, V.M. Ruiz-Valdiviezo, and F.A. Gutierrez-Miceli. 2017. Vermicompost and vermiwash minimized the influence of salinity stress on growth parameters in potato plants. Compost Science & Utilization. 12: 1-7.
Rykaczewska, K. 2013. The impact of high temperature during growing season on potato cultivars with different response to environmental stresses. American Journal of Plant Sciences. 4: 2386–2393.
Schlegel, H.G. 1956. Die verwertung organischer sauren durch chlorella in lincht. Planta. 47: 510-515.
Shibli, R.A., M. Kushad, G.G. Yousef, and M. Lila, 2007. Physiological and biochemical responses of tomato micro shoots to induced salinity stress with associated ethylene accumulation. Plant Growth Regulation. 51: 159-169.
Shimazaki, T., T. Endo, M. Kasuga, K. Yamaguchi-Shinozaki, K.N. Watanabe, and A. Kikuchi. 2017. Evaluation of the yield of abiotic-stress-tolerant tDREB1A transgenic potato under saline conditions in advance of field trials. Breeding Science. 66: 703–710.
Simersky, R., O. Nova´, K.D.A. Morris, V. Pouzar, and M. Strnad .2009. Identification and quantification of several mammalian steroid hormones in plants by UPLC–MS/MS. Journal of Plant Growth Regulation. 28: 125–136.
Sudhersan, C., Jibi Manuel, J. Ashkanani, and A. Al-Ajeel. 2012. In vitro screening of potato cultivars for salinity tolerance. American-Eurasian Journal of Sustainable Agriculture. 6(4): 344-348.
Uranbey, S., D. Köm, Akdoğan, H.A.A. Ahmed, N. Koçak, and M.E. Kara. 2017. The effects of salt stress on expression of Asg1 gene related stomatal resistance in potato. Mediterranean Agricultural Sciences. 30(3): 235-238.
Velikova, V., I. Yordanov, and A. Edreva .2000. Oxidative stress and some antioxidant systems in acid rain treated bean plants. Protective role of exogenous polyamines. Plant Science. 151: 59-66.
Wang, L., Y. Liu, D. Li, S. Feng, J. Yang, J. Zhang, J. Zhang, Wang, and Y. Gan. 2017. Improving salt tolerance in potato through overexpression of AtHKT1 gene. BMC Plant Biology. 9(357): 1-57.
Zaman, S.M., M. Ali, A. Muhammad, and I. Hussain. 2015. In vitro screening of salt tolerance in potato (Solanum tuberosum L.) varieties. Sarhad Journal of Agriculture. 31(2): 106-113.