بررسی عملکرد دانه و برخی خصوصیات بیـوشیمیایی پنج رقم نخود (.Cicer arietinum L) تحت تنش خشکی در منطقه کرمانشاه
محورهای موضوعی : اکوفیزیولوژی گیاهان زراعیسیدمحمد ناصح حسینی 1 , محسن سعیدی 2 , سیروس منصوری فر 3
1 - فرهیختهی کارشناسی ارشد، گروه زراعت و اصلاح نباتات، پردیس کشاورزی و منابع طبیعی، دانشگاه رازی، کرمانشاه، ایران
2 - دانشیار گروه زراعت و اصلاح نباتات، دانشگاه رازی، کرمانشاه، ایران
3 - دانشیار گروه کشاورزی، دانشگاه پیام نور البرز، کرج، ایران
کلید واژه: عملکرد, کلروفیل, تنش کم آبی, آنزیمهای آنتیاکسیدان, کاروتنوئیدها,
چکیده مقاله :
به منظور بررسی تغییرات بیوشیمیایی ناشی از تنش کم آبی در نخود، آزمایشی به صورت کرت های خرد شده و در قالب طرح پایه بلوک های کامل تصادفی با سه تکرار در دانشگاه رازی کرمانشاه اجرا شد. عامل اصلی تنش کم آبی با سه سطح شامل: 1- تنش کم آبی ابتدای گلدهی تا رسیدگی فیزیولوژیک، 2- تنش کم آبی از ابتدای غلاف دهی تا رسیدگی فیزیولوژیک، 3- آبیاری مطلوب و عامل فرعی پنج رقم نخود شامل آرمان، آزاد، بیونیج، هاشم و ILC482 بودند. بر اساس نتایج، تنش کم آبی در هر دو سطح موجب کاهش معنی دار محتوی کلروفیل ها و کاروتنوئیدها و افزایش معنی دار فعالیت آنزیم های آنتی اکسیدان نظیر: پراکسیداز، کاتالاز و سوپراکسیددیسموتاز در برگ ها در مقایسه با تیمار شاهد شد. بنابراین، بین فعالیت آنزی م های آنتی اکسیدان با میزان آب قابل استفاده در خاک رابطه ی منفی وجود داشت و فعالیت آنها با افزایش شدت تنش کم آبی به طور معنی داری افزایش یافت. عملکرد دانه به طور معنی داری تحت تأثیر کمبود آب قرار گرفت و تیمار تنش کم آبی از زمان ابتدای گلدهی تا رسیدگی، بیشتر از دیگر تیمار تنش کم آبی (ابتدای غلاف دهی تا رسیدگی) موجب کاهش عملکرد دانه، به ترتیب 36 و 15 درصد، نسبت به شرایط بدون تنش شد. در بین ارقام مورد بررسی، تحت تنش کم آبی از شروع گلدهی، رقم ILC482 با عملکرد دانه به میزان 715 (کیلوگرم در هکتار) و در شرایط عدم وجود تنش، رقم آرمان با عملکرد دانه به مقدار 1355 (کیلوگرم در هکتار) عملکرد مناسب تری از خود نشان دادند. ILC482، آزاد و بیونیج به عنوان ارقام با عملکرد بالاتر در شرایط وقوع تنش کم آبی در هر دو تیمار تنش کم آبی، همچنین دارای محتوی بیشتر رنگیزه ها و سرعت فعالیت بیشتر آنزیم های آنتی اکسیدان در برگ های خود بودند. نتایج حاصله حاکی از اثرات مثبت محتوای رنگیزه های گیاهی و فعالیت آنزیم های آنتی اکسیدان جهت افزایش توان تحملی نخود برای تداوم رشد و حفظ عملکرد قابل قبول در شرایط تنش خشکی بود.
To study the effect of water deficit on biochemical changes in chickpea, a split-plot experiment based on complete block design with three replications was carried out at the Research Farm of Razi University in Iran. Moisture regimes with three levels, were: 1) irrigation cut off from beginning of flowering till maturity, 2) irrigation cut off from beginning of podding till maturity and 3) irrigating plants at all stages of growth (control) assigned to the main-plots and five chickpea cultivars: Arman, Azad, Bivanij, Hashem and ILC482 to the sub-plots. Based on the results, two levels of water deficit resulted in significant reduction in the chlorophylls and carotenoids concentration and significant increase in the activity of antioxidant enzymes such as: catalase, peroxidase and super oxide dismutase of leaves, in comparison with control. Therefore, there was a negative correlation between the activity of antioxidant enzymes and the amount of available water in soil, and their activity increased with increasing the severity of water stress. Seed yield was significantly affected by water deficit. In comparison of control treatment, irrigation cut off from the beginning of flowering up to maturity compared to irrigation cut off from podding up to maturity resulted in more reduction in seed yield (36 and 15% respectively). ILC482 cultivar with seed yield of 715 kg.ha-1 under irrigation cut off from the beginning of flowering up to maturity and Arman cultivar with a seed yield of 1355 kg.ha-1 under irrigation cut off from podding up to maturity produced highest grain yield. High yield cultivars under two levels of water deficit including: ILC482, Azad and Bivanij also had the highest photosynthetic pigments concentration and highest antioxidant activity in their leaves. The results also indicated a positive relationship between the antioxidant enzymes activities and photosynthetic pigment concentrations in chickpea which may help to increase growth and yield of chickpea under drought stress conditions.
· Abbaslu, L., S.A.R. Kazemini, M. Edalat, and A. Dadkhodaee. 2014. Effect of drought stress and planting arrangement on some physiological and biochemical characteristics of two chickpea cultivars. Journal of Agricultural Improvment. 16(4): 933-943. (In Persian).
· Abhari, A., E. Azizi, and B. Hareth-Abadi. 2017. Effect of super absorbent on yield and yield components of chickpea under drought stress conditions of the end of season. Crop Production Publication. 10(1): 191-202. (In Persian).
· Akashi, K., C. Miyake, and A. Yakota. 2001. Citrulline, a novel compatible solute in drought tolerant wild watermelon leaves, is an efficient hydroxyl radical scavenger.FEBS Lett. 508: 438-442.
· Alscher, R.G., N. Erturk, and L.S. Heath. 2002. Role of superoxidedismotase (SOD) in controlling oxidative stress in plant. Journal of Experimental Botany. 153: 1331-1341.
· Anonymous. 2016. FAO. http://faostat.fao.org/site/339/default.aspx.
· Awari, V.R., U.S. Dalvi, P.K. Lokhande, V.Y. Pawar, S.N. Mate, R.M. Naik, and L.B. Mhase. 2017. Physiological and biochemical basis for moisture stress tolerance in chickpea under pot study. International Journal of Current Microbiology and Applied Sciences. 6(5): 1247-1259.
· Azpilicueta, C.E., M.P. Benavides, M.L. Tomaro, and S.M. Gallego. 2007. Mechanism of CATA3 induction by cadmium in sunflower leaves. Plant Physiology and Biochemistry. 45: 589-595.
· Beauchamp, C., and I. Fridovich. 1971. Superoxide dismutase improved assays andan assay predictable to acrylamide gels. Annals of Biochemistry. 44: 276–287.
· Behra, R.K., P.C. Mishra, and N.K. Choudhury. 2002. High irradiance and water stress induce alterations in pigment composition and chloroplast activities of primary wheat leaves. Journal Plant Physiology. 159: 967-973.
· Blokhina, O., E. Virolainen, and K.V. Fagerstedt. 2003. Antioxidants, oxidative damage and oxygen deprivation stress. Annual Review of Botany. 91: 179-194.
· Chance, B., and A.C. Maehly. 1995. Assay of catalase and peroxidase. PP. 764-765 In: S.P. Culowic, and N.O. Kaplan (eds). Methods in enzymology Vol. 2. Academic Press. Inc. New York.
· Choudhury, F.K., R.M. Rivero, E. Blumwald, and R. Mittler. 2017. Reactive oxygen species, abiotic stress and stress combination. Plant Journal. 90(5): 856- 867.· Dadkhah, N., A. Ebadi, G. Parmoon, E. Gholipoori, and S. Jahanbakhsh. 2014. Effect of spraying zinc on photosynthetic pigments and grain yield of chickpea under level different irrigation. Iranian Agriculture Drought Journal. 2(2): 141-161. (In Persian).
· Edreva, A. 2005. Generation and scavenging of reactive oxygen species in chloroplasts a submolecular approach. Agriculture Ecosystems and Environment. 106: 119-133.
· Fang, X., N.C. Turner, G. Yan, F. Li, and K.H.M. Siddique. 2010. Flower numbers, pod production, pollen viability, and pistil function are reduced and flower and pod abortion increased in chickpea (Cicer arietinum L.) under terminal drought. Journal of Experimental Botany. 61: 335-345.
· Farooq, M., A. Wahid, N. Kobayashi, D. Fujita, and S.M.A. Basra. 2009. Plant drought stress: effects, mechanisms and management. Agronomy for Sustainable Development. 29: 185–212.
· Farshadfar, E., and J. Javadiniya. 2011. Evaluation of chickpea (Cicer arietinum L.) genotypes for drought tolerance. Journal to Breed of Seed and Seedling. 27-1(4): 517- 537. (In Persian).
· Flexas, J., J. Bota, F. Loreto, G. Cornic, and T.D. Sharkey. 2004. Diffusive and metabolic limitations to photosynthesis under drought and salinity in C3 plants. Plant Biology. 6: 1-11.
· Ghasemi-Golazani, K., S. Mohamadi, P. Rahem-Zadeh, and M. Moghadam. 2013. Quantitative connection between density and yield of three chickpea cultivar on different planting dates. Journal of Plant Physiology and Breeding. 7: 59-73.
· Ghorbanli, M., G.R. Bakhshi-Khaniki, S. Salimi-Elizei, and M. Hedayati. 2010. Effect of water deficit and its interaction with ascorbate on proline, soluble sugars, catalase and glutathione peroxidise amounts in Nigella sativa L. Iranian Journal of Medicinal and Aromatic Plants. 26(4): 46-476. (In Persian).
Gregersen, P.L., and P.B. Holm. 2007. Transcriptome analysis of senescence in the flag leaf of wheat. Plant Biotechnology. 5: 192-206.
· Hossinzadeh, S.R., A. Salimi, A. Ganjeali, and R. Ahmadpour. 2015. Effect of foliar application of methanol on biochemical characteristics and antioxidant enzyme activity of chickpea under drought stress. Journal of Plant Physiology and Biochemistry. 1(1): 17-30. (In Persian).
Izanloo, A., A.G. Condon, P. Langridge, M. Tester, and T. Schnurbusch. 2008. Different mechanisms of adaptation to cyclic water stress in two South Australian bread wheat cultivars. Journal of Experimental Botany. 59(12): 3327-3346.
· Jabari, F., A. Ahmadi, K. Poustini, and H. Alizadeh. 2006. Evaluation of some antioxidant enzyme effects on chlorophyll and cell membrane in drought susceptible and tolerant wheat varieties. Iranian Journal of Agricultural Science. 37: 307-316. (In Persian).
· Kamrava, S., N. Babaeianjolodar, and N. Bagheri. 2017. Evaloation of drought stress on chlorophyll and proline traits in soybean genotypes. Journal of Crop Breeding. 9(23): 95-104. (In Persian).
· Kanuni, H., H. Kazemi, M. Moghaddam, and M.R. Neyshburi. 2002. Selection of chickpea (Cicer arietinum L.) lines for drought resistance. Journal of Agricultural Science. 12(2): 109-121. (In Persian).
· Kashiwagi, J., L. Krishnamurthy, R. Purushothaman, H.D. Upadhyaya, P.M. Gaur, C.L.L. Gowda, and R.K. Varshney. 2015. Scope for improvement of yield under drought through the root traits in chickpea (Cicer arietinum L.). Field Crops Research. 170: 47-54.
· Khan, N., A. Bano, M.A. Rahman, B. Rathinasabapathi, and M.A. Babar. 2018. UPLC-HRMS-based untargeted metabolic profiling reveals changes in chickpea metabolome following long-term drought stress. Plant, Cell and Environment. 42(1): 115-132.
· Kouchaki, A., and M. Bannayan-Aval. 1994. Yield physiology in crops. Jahad Daneshgahi Mashhad Publisher. 261 pp. (In Persian).
· Krishnamurthy, A., and B. Rathinasabapathi. 2013. Oxidative stress tolerance in plants: novel interplay between auxin and reactive oxygen species signaling. Plant Signaling and Behavior. 8(5): 257-261.
· Lichtenthaler, H., and A.R. Wellburn. 1983. Determination of total carotenoids and chlorophyll a and chlorophyll b leaf extracts in different solvents. Biochemical Society Transactions. 603: 591-592.
· Mafakheri A., A. Sio-Semardeh, B. Bahramnejad, and Y. Sohrabi. 2010. Effect of drought stress on yield, proline and chlorophyll contents in three chickpea cultivars. Australian Journal of Crop Science. 4(8): 580-585.
· Mafakheri, A., A. Sio-Semardeh, B. Bahramnejad, P.C. Struik, and Y. Sohrabi. 2011. Effect of drought stress and subsequent recovery on protein, carbohydrate contents, catalase, and peroxidase activities in three chickpea (Cicer arietinum) cultivars. Australian Journal of Crop Science. 5(10): 1255-1260.
Maiti, R.K., S. Moreno-Limon, and P. Wesche-Ebeling. 2000. Responses of some crops to various abiotic stress factors and its physiological and biochemical basis of resistances. Agricultural Reviews. 21: 155-167.
· Majnoon-Hosseini, N., H. Mohammadi, K. Poustini, and H. Zeinaly-khanghah. 2003. Effect of plant density on agronomic characteristics, chlorophyll content and stem remobilization percentage in chickpea cultivars (Cicer arietinum L.). Iranian Journal Agriculture Science. 34(4): 1011-1019. (In Persian).
· Makarian, H., H. Shojaei, A. Damavandi, A. Nasiri-Dehsorkhi, and A. Akhyani. 2017. The effect of foliar application of zinc oxide in common and nanoparticles forms on some growth and quality traits of mungbean (Vigna radiata L.) under drought stress conditions. Iranian Journal of Pulses Research. 8(2): 166-180. (In Persian).
· Maleky, A., A. Heidary-Moghaddam, S.A. Siyadat, and A. Tahmasebi. 2011. Effect of supplemental irrigation on yield, yield components and seed protein percentage of three chickpea cultivars in Ilam. Journal of Crop Ecophysiology. 19(5): 65-78. (In Persian).
· Menconi, M., C.L.M. Sgherri, C. Pinzino, and F. Navari-Izzo. 1995. Activated oxygen production and detoxification in wheat plants subjected to a water deficit programme. Journal of Experimental Botany. 46:1123-1130.
· Milone, M.T., C. Sgherri, H. Clijters, and F. Navari-Izzo. 2003. Antioxidative responses of wheat treated with realistic concentrations of cadmium. Environment and Experimental Botany. 50: 265-273.
· Mirzaee, M., A. Moieni, and F. Ghanati. 2013. Effects of drought stress on the lipid peroxidation and antioxidant enzyme activity in two canola (Brassica napus L.) cultivars. Journal of Agriculture and Science Technology. 15: 593-602.
· Mirzavand, M., KH. Azizi, M. Abdali, and A. Esmaeili. 2011. Effect of some agricultural techniques (Planting arrangement and supplementary irrigation) on chickpea growth indices. Journal of Crop Ecophysiology. 2(3): 63-73. (In Persian).
· Mittler, R., S. Vanderauwera, N. Suzuki, G. Miller, V.B. Tognetti, K. Vandepoele, M. Gollery, V. Shulaev, and F. Van Breusegem. 2011. ROS signaling: the new wave?. Trends Plant Science. 16(6): 300-309.
· Mohammadi, A., D. Habibi, M. Rohami, and S. Mafakheri. 2011. Effect of drought stress on antioxidant enzymes activity of some chickpea cultivars. American-Eurasian Journal of Agriculture and Environmental Science. 11(6): 782-785.
· Mohammadi, M., M. Roozrokh, and R. Talebi. 2016. Effect of supplemental irrigation and iron foliar application on Chickpea genotypes in Kermanshah. Journal of Crop Ecophysiology. 27: 103-113. (In Persian).
· Nasr-Esfahani, M. 2013. Effect of dry stress on growth and antioxidant system in three chickpea (Cicer arietinum L.) cultivars. Journal of Plant Biology. 15: 111-124.
· Nemati, A., M. Rafieealhusseini, and A. Danesh-Shahraki. 2016. Effect of livestock manure and bacterial inoculation on physiological indices, yield and yield components of chickpea (Cicer arietinum L.) under drought stress. Environmental Stresses in Crop Sciences. 9(4): 339-351. (In Persian).
· Oberoi, H.K., A.K. Gupeta, S. Kaur, and I. Singh. 2014. Stage specific upregulation of antioxidant defence system in leaves for regulating drought tolerance in chickpea. Journal of Applied Natural Science. 6(2): 326-337.
· Patel, P.K., A. Hemantaranjan, B.K. Sarma, and R. Singh. 2011. Growth and antioxidant system under drought stress in chickpea (Cicer arietinum) as sustained by salicylic acid. Journal of Stress Physiology and Biochemistry. 7: 130-144.
· Patel, P.K., and A. Hemantaranjan. 2012. Antioxidant defence system in chickpea (Cicer arietinum L.) influence by drought stress implemented at pre and post anthesis stage. American Journal of Plant Physiology. 7(4): 164-173.
· Sarvajeet, S.G., and T. Narendra. 2010. Reactive oxygen species and antioxidant machinery in a biotic stress tolerance in crop plants. Plant Physiology and Biochemistry. 3: 1-22.
· Shigeoka, S., T. Ishikawa, M. Tamoi, Y. Miyagawa, T. Takeda, Y. Yabuta, and K. Yoshimura. 2002. Regulation and function of ascorbate peroxidase isoenzymes. Journal of Experimental Botany. 53: 1305-1319.
· Shuryabi, M., A. Gangeali, and P. Abrishamchi. 2012. Effect of salicylic acid on the activity of enzyme and antioxidant compounds in chickpea cultivars at facing drought stress. Journal of Environmental Tensions in Crop Sciences. 1(5): 41-54. (In Persian).
· Singh, D.P. 2003. Oxidative stress in stress physiology. New Age International Limited. New Dehli. 41 pp.
· Singh, K.B., G. Bejiga, and R.S. Malhorta. 1990. Assosiations of some characters with seed yield in chickpea collection. Euphytica. 49(1): 83-88.
· Sinha, A.K. 1972. Colorimetric assay of catalase. Analytical Biochemistry. 47(2): 389-394.
· Sirousmehr, A., J. Bardel, and S. Mohammadi. 2014. Changes of Germination Properties, Photosynthetic Pigments and antioxidant enzymes activity of safflower as affected by drought and salinity stresses. Journal of Crop Ecophysiology. 8(4): 517-534. (In Persian).· Tale-Ahmad, S., and R. Hadad. 2010. Effect of silicon on antioxidant enzymes activities and osmotic adjustment contents in two bread wheat genotypes under drought stress conditions. Journal of Seed and Seedling Crop. 26-2(2): 207-225. (In Persian).
· Talebi R., M.H. Ensafi, N. Baghebani, E. Karami, and K. Mohammadi. 2013. Physiological responses of chickpea (Cicer arietinum) genotypes to drought stress. Environmental and Experimental Biology. 11: 9-15.
Tanaka, A., and R. Tanaka. 2006. Chlorophyll metabolism. Plant Biology. 9: 248- 255.
· Thaman, M., A. Sepehri, G. Ahmadvand, and S.H. Sabbaghpour. 2007. Effect of irrigation in pod formation and seed filling on growth and yield of five chickpea cultivar. Journal of Agricultural Research, Water, Soil and Plant in Agriculture. 7(1): 41-59. (In Persian).
· Yadav, R.S., C.T. Hash, F.R. Bidinger, K.M. Devos, and C.J. Howarth. 2004. Genomic regions associated with grain yield and aspects of post flowering drought tolerance in pearl millet across environments and tester background. Euphytica. 136: 265-277.
Yarnia, M. 2015. The effect of water deficit stress on osmotic metabolites and antioxidant system and grain and oil yield of amaranth CV. koniz. Journal of Crop Ecophysiology. 8(4): 499-516. (In Persian).
· Abbaslu, L., S.A.R. Kazemini, M. Edalat, and A. Dadkhodaee. 2014. Effect of drought stress and planting arrangement on some physiological and biochemical characteristics of two chickpea cultivars. Journal of Agricultural Improvment. 16(4): 933-943. (In Persian).
· Abhari, A., E. Azizi, and B. Hareth-Abadi. 2017. Effect of super absorbent on yield and yield components of chickpea under drought stress conditions of the end of season. Crop Production Publication. 10(1): 191-202. (In Persian).
· Akashi, K., C. Miyake, and A. Yakota. 2001. Citrulline, a novel compatible solute in drought tolerant wild watermelon leaves, is an efficient hydroxyl radical scavenger.FEBS Lett. 508: 438-442.
· Alscher, R.G., N. Erturk, and L.S. Heath. 2002. Role of superoxidedismotase (SOD) in controlling oxidative stress in plant. Journal of Experimental Botany. 153: 1331-1341.
· Anonymous. 2016. FAO. http://faostat.fao.org/site/339/default.aspx.
· Awari, V.R., U.S. Dalvi, P.K. Lokhande, V.Y. Pawar, S.N. Mate, R.M. Naik, and L.B. Mhase. 2017. Physiological and biochemical basis for moisture stress tolerance in chickpea under pot study. International Journal of Current Microbiology and Applied Sciences. 6(5): 1247-1259.
· Azpilicueta, C.E., M.P. Benavides, M.L. Tomaro, and S.M. Gallego. 2007. Mechanism of CATA3 induction by cadmium in sunflower leaves. Plant Physiology and Biochemistry. 45: 589-595.
· Beauchamp, C., and I. Fridovich. 1971. Superoxide dismutase improved assays andan assay predictable to acrylamide gels. Annals of Biochemistry. 44: 276–287.
· Behra, R.K., P.C. Mishra, and N.K. Choudhury. 2002. High irradiance and water stress induce alterations in pigment composition and chloroplast activities of primary wheat leaves. Journal Plant Physiology. 159: 967-973.
· Blokhina, O., E. Virolainen, and K.V. Fagerstedt. 2003. Antioxidants, oxidative damage and oxygen deprivation stress. Annual Review of Botany. 91: 179-194.
· Chance, B., and A.C. Maehly. 1995. Assay of catalase and peroxidase. PP. 764-765 In: S.P. Culowic, and N.O. Kaplan (eds). Methods in enzymology Vol. 2. Academic Press. Inc. New York.
· Choudhury, F.K., R.M. Rivero, E. Blumwald, and R. Mittler. 2017. Reactive oxygen species, abiotic stress and stress combination. Plant Journal. 90(5): 856- 867.· Dadkhah, N., A. Ebadi, G. Parmoon, E. Gholipoori, and S. Jahanbakhsh. 2014. Effect of spraying zinc on photosynthetic pigments and grain yield of chickpea under level different irrigation. Iranian Agriculture Drought Journal. 2(2): 141-161. (In Persian).
· Edreva, A. 2005. Generation and scavenging of reactive oxygen species in chloroplasts a submolecular approach. Agriculture Ecosystems and Environment. 106: 119-133.
· Fang, X., N.C. Turner, G. Yan, F. Li, and K.H.M. Siddique. 2010. Flower numbers, pod production, pollen viability, and pistil function are reduced and flower and pod abortion increased in chickpea (Cicer arietinum L.) under terminal drought. Journal of Experimental Botany. 61: 335-345.
· Farooq, M., A. Wahid, N. Kobayashi, D. Fujita, and S.M.A. Basra. 2009. Plant drought stress: effects, mechanisms and management. Agronomy for Sustainable Development. 29: 185–212.
· Farshadfar, E., and J. Javadiniya. 2011. Evaluation of chickpea (Cicer arietinum L.) genotypes for drought tolerance. Journal to Breed of Seed and Seedling. 27-1(4): 517- 537. (In Persian).
· Flexas, J., J. Bota, F. Loreto, G. Cornic, and T.D. Sharkey. 2004. Diffusive and metabolic limitations to photosynthesis under drought and salinity in C3 plants. Plant Biology. 6: 1-11.
· Ghasemi-Golazani, K., S. Mohamadi, P. Rahem-Zadeh, and M. Moghadam. 2013. Quantitative connection between density and yield of three chickpea cultivar on different planting dates. Journal of Plant Physiology and Breeding. 7: 59-73.
· Ghorbanli, M., G.R. Bakhshi-Khaniki, S. Salimi-Elizei, and M. Hedayati. 2010. Effect of water deficit and its interaction with ascorbate on proline, soluble sugars, catalase and glutathione peroxidise amounts in Nigella sativa L. Iranian Journal of Medicinal and Aromatic Plants. 26(4): 46-476. (In Persian).
Gregersen, P.L., and P.B. Holm. 2007. Transcriptome analysis of senescence in the flag leaf of wheat. Plant Biotechnology. 5: 192-206.
· Hossinzadeh, S.R., A. Salimi, A. Ganjeali, and R. Ahmadpour. 2015. Effect of foliar application of methanol on biochemical characteristics and antioxidant enzyme activity of chickpea under drought stress. Journal of Plant Physiology and Biochemistry. 1(1): 17-30. (In Persian).
Izanloo, A., A.G. Condon, P. Langridge, M. Tester, and T. Schnurbusch. 2008. Different mechanisms of adaptation to cyclic water stress in two South Australian bread wheat cultivars. Journal of Experimental Botany. 59(12): 3327-3346.
· Jabari, F., A. Ahmadi, K. Poustini, and H. Alizadeh. 2006. Evaluation of some antioxidant enzyme effects on chlorophyll and cell membrane in drought susceptible and tolerant wheat varieties. Iranian Journal of Agricultural Science. 37: 307-316. (In Persian).
· Kamrava, S., N. Babaeianjolodar, and N. Bagheri. 2017. Evaloation of drought stress on chlorophyll and proline traits in soybean genotypes. Journal of Crop Breeding. 9(23): 95-104. (In Persian).
· Kanuni, H., H. Kazemi, M. Moghaddam, and M.R. Neyshburi. 2002. Selection of chickpea (Cicer arietinum L.) lines for drought resistance. Journal of Agricultural Science. 12(2): 109-121. (In Persian).
· Kashiwagi, J., L. Krishnamurthy, R. Purushothaman, H.D. Upadhyaya, P.M. Gaur, C.L.L. Gowda, and R.K. Varshney. 2015. Scope for improvement of yield under drought through the root traits in chickpea (Cicer arietinum L.). Field Crops Research. 170: 47-54.
· Khan, N., A. Bano, M.A. Rahman, B. Rathinasabapathi, and M.A. Babar. 2018. UPLC-HRMS-based untargeted metabolic profiling reveals changes in chickpea metabolome following long-term drought stress. Plant, Cell and Environment. 42(1): 115-132.
· Kouchaki, A., and M. Bannayan-Aval. 1994. Yield physiology in crops. Jahad Daneshgahi Mashhad Publisher. 261 pp. (In Persian).
· Krishnamurthy, A., and B. Rathinasabapathi. 2013. Oxidative stress tolerance in plants: novel interplay between auxin and reactive oxygen species signaling. Plant Signaling and Behavior. 8(5): 257-261.
· Lichtenthaler, H., and A.R. Wellburn. 1983. Determination of total carotenoids and chlorophyll a and chlorophyll b leaf extracts in different solvents. Biochemical Society Transactions. 603: 591-592.
· Mafakheri A., A. Sio-Semardeh, B. Bahramnejad, and Y. Sohrabi. 2010. Effect of drought stress on yield, proline and chlorophyll contents in three chickpea cultivars. Australian Journal of Crop Science. 4(8): 580-585.
· Mafakheri, A., A. Sio-Semardeh, B. Bahramnejad, P.C. Struik, and Y. Sohrabi. 2011. Effect of drought stress and subsequent recovery on protein, carbohydrate contents, catalase, and peroxidase activities in three chickpea (Cicer arietinum) cultivars. Australian Journal of Crop Science. 5(10): 1255-1260.
Maiti, R.K., S. Moreno-Limon, and P. Wesche-Ebeling. 2000. Responses of some crops to various abiotic stress factors and its physiological and biochemical basis of resistances. Agricultural Reviews. 21: 155-167.
· Majnoon-Hosseini, N., H. Mohammadi, K. Poustini, and H. Zeinaly-khanghah. 2003. Effect of plant density on agronomic characteristics, chlorophyll content and stem remobilization percentage in chickpea cultivars (Cicer arietinum L.). Iranian Journal Agriculture Science. 34(4): 1011-1019. (In Persian).
· Makarian, H., H. Shojaei, A. Damavandi, A. Nasiri-Dehsorkhi, and A. Akhyani. 2017. The effect of foliar application of zinc oxide in common and nanoparticles forms on some growth and quality traits of mungbean (Vigna radiata L.) under drought stress conditions. Iranian Journal of Pulses Research. 8(2): 166-180. (In Persian).
· Maleky, A., A. Heidary-Moghaddam, S.A. Siyadat, and A. Tahmasebi. 2011. Effect of supplemental irrigation on yield, yield components and seed protein percentage of three chickpea cultivars in Ilam. Journal of Crop Ecophysiology. 19(5): 65-78. (In Persian).
· Menconi, M., C.L.M. Sgherri, C. Pinzino, and F. Navari-Izzo. 1995. Activated oxygen production and detoxification in wheat plants subjected to a water deficit programme. Journal of Experimental Botany. 46:1123-1130.
· Milone, M.T., C. Sgherri, H. Clijters, and F. Navari-Izzo. 2003. Antioxidative responses of wheat treated with realistic concentrations of cadmium. Environment and Experimental Botany. 50: 265-273.
· Mirzaee, M., A. Moieni, and F. Ghanati. 2013. Effects of drought stress on the lipid peroxidation and antioxidant enzyme activity in two canola (Brassica napus L.) cultivars. Journal of Agriculture and Science Technology. 15: 593-602.
· Mirzavand, M., KH. Azizi, M. Abdali, and A. Esmaeili. 2011. Effect of some agricultural techniques (Planting arrangement and supplementary irrigation) on chickpea growth indices. Journal of Crop Ecophysiology. 2(3): 63-73. (In Persian).
· Mittler, R., S. Vanderauwera, N. Suzuki, G. Miller, V.B. Tognetti, K. Vandepoele, M. Gollery, V. Shulaev, and F. Van Breusegem. 2011. ROS signaling: the new wave?. Trends Plant Science. 16(6): 300-309.
· Mohammadi, A., D. Habibi, M. Rohami, and S. Mafakheri. 2011. Effect of drought stress on antioxidant enzymes activity of some chickpea cultivars. American-Eurasian Journal of Agriculture and Environmental Science. 11(6): 782-785.
· Mohammadi, M., M. Roozrokh, and R. Talebi. 2016. Effect of supplemental irrigation and iron foliar application on Chickpea genotypes in Kermanshah. Journal of Crop Ecophysiology. 27: 103-113. (In Persian).
· Nasr-Esfahani, M. 2013. Effect of dry stress on growth and antioxidant system in three chickpea (Cicer arietinum L.) cultivars. Journal of Plant Biology. 15: 111-124.
· Nemati, A., M. Rafieealhusseini, and A. Danesh-Shahraki. 2016. Effect of livestock manure and bacterial inoculation on physiological indices, yield and yield components of chickpea (Cicer arietinum L.) under drought stress. Environmental Stresses in Crop Sciences. 9(4): 339-351. (In Persian).
· Oberoi, H.K., A.K. Gupeta, S. Kaur, and I. Singh. 2014. Stage specific upregulation of antioxidant defence system in leaves for regulating drought tolerance in chickpea. Journal of Applied Natural Science. 6(2): 326-337.
· Patel, P.K., A. Hemantaranjan, B.K. Sarma, and R. Singh. 2011. Growth and antioxidant system under drought stress in chickpea (Cicer arietinum) as sustained by salicylic acid. Journal of Stress Physiology and Biochemistry. 7: 130-144.
· Patel, P.K., and A. Hemantaranjan. 2012. Antioxidant defence system in chickpea (Cicer arietinum L.) influence by drought stress implemented at pre and post anthesis stage. American Journal of Plant Physiology. 7(4): 164-173.
· Sarvajeet, S.G., and T. Narendra. 2010. Reactive oxygen species and antioxidant machinery in a biotic stress tolerance in crop plants. Plant Physiology and Biochemistry. 3: 1-22.
· Shigeoka, S., T. Ishikawa, M. Tamoi, Y. Miyagawa, T. Takeda, Y. Yabuta, and K. Yoshimura. 2002. Regulation and function of ascorbate peroxidase isoenzymes. Journal of Experimental Botany. 53: 1305-1319.
· Shuryabi, M., A. Gangeali, and P. Abrishamchi. 2012. Effect of salicylic acid on the activity of enzyme and antioxidant compounds in chickpea cultivars at facing drought stress. Journal of Environmental Tensions in Crop Sciences. 1(5): 41-54. (In Persian).
· Singh, D.P. 2003. Oxidative stress in stress physiology. New Age International Limited. New Dehli. 41 pp.
· Singh, K.B., G. Bejiga, and R.S. Malhorta. 1990. Assosiations of some characters with seed yield in chickpea collection. Euphytica. 49(1): 83-88.
· Sinha, A.K. 1972. Colorimetric assay of catalase. Analytical Biochemistry. 47(2): 389-394.
· Sirousmehr, A., J. Bardel, and S. Mohammadi. 2014. Changes of Germination Properties, Photosynthetic Pigments and antioxidant enzymes activity of safflower as affected by drought and salinity stresses. Journal of Crop Ecophysiology. 8(4): 517-534. (In Persian).· Tale-Ahmad, S., and R. Hadad. 2010. Effect of silicon on antioxidant enzymes activities and osmotic adjustment contents in two bread wheat genotypes under drought stress conditions. Journal of Seed and Seedling Crop. 26-2(2): 207-225. (In Persian).
· Talebi R., M.H. Ensafi, N. Baghebani, E. Karami, and K. Mohammadi. 2013. Physiological responses of chickpea (Cicer arietinum) genotypes to drought stress. Environmental and Experimental Biology. 11: 9-15.
Tanaka, A., and R. Tanaka. 2006. Chlorophyll metabolism. Plant Biology. 9: 248- 255.
· Thaman, M., A. Sepehri, G. Ahmadvand, and S.H. Sabbaghpour. 2007. Effect of irrigation in pod formation and seed filling on growth and yield of five chickpea cultivar. Journal of Agricultural Research, Water, Soil and Plant in Agriculture. 7(1): 41-59. (In Persian).
· Yadav, R.S., C.T. Hash, F.R. Bidinger, K.M. Devos, and C.J. Howarth. 2004. Genomic regions associated with grain yield and aspects of post flowering drought tolerance in pearl millet across environments and tester background. Euphytica. 136: 265-277.