Document Type : Original Article

Authors

1 PhD student in Agricultural Biotechnology, Faculty of Agriculture, Ferdowsi University of Mashhad, Iran

2 Department of Biotechnology and Plant Breeding, Faculty of Agriculture, Ferdowsi University of Mashhad, Iran

3 Department of Ornamental Plants Biotechnology, Academic Center of Education, Culture and Research of Mashhad, Iran

Abstract

Introduction
Plants are exposed to various environmental stresses during growth and development under natural and agricultural conditions. Drought is one of the most severe abiotic stresses that greatly affects plant yield. Metabolic changes under stress cause the morphological and physiological changes in the plant which may eventually lead to reduced yield. Barley is one of the most important cereals which a large part of the human population in many parts of the world are dependent on it as a source of food and animal feed.
Materials and methods
In order to investigate the effects of drought stress on some morphological and physiological traits in barley as one of the most important crop, this experiment was carried out in the greenhouse of the agriculture faculty, Ferdowsi University of Mashhad in 2019 in a factorial based on a randomized complete block design with three replications. Experimental factors were included: different levels of drought stress including 100% of field capacity (control or no stress), 80, 60 and 40% of field capacity and two barley cultivars containing semi-sensitive (Fajr30) and drought tolerant (Dasht). First, the seeds of both cultivars were planted in trays filled with coco peat, perlite and sand, and after two weeks, the seedlings were transferred to pots filled with garden soil. The pots were watered daily and after one month (4 to 6 leaf stage) until the end of the growing season, they were subjected to drought stress treatments. During the vegetative stage, morphological traits such as plant height (from crown to end of plant), stem diameter, number of leaves, number of tillers, leaf area and number of stomata below and above the leaf were examined using common methods. Physiological traits including proline, soluble sugars, photosynthetic pigments and antioxidant enzymes (catalase and peroxidase) were also measured. Analysis of variance was performed using JMP statistical software version 8 and the mean of treatments was compared using LSD test at the level of 5%.
Results and discussion
The results of this experiment showed that the effect of cultivar and drought stress were significant for many of the studied traits. The highest plant height, stem diameter, number of leaves and tillers and leaf area index were observed in plants without stress (100% FC), which was significant compared to other stress level. Also, the comparison of the two cultivars showed that Dasht, as a drought tolerant cultivar, has more leaves and tillers but less leaf area and stomata than the semi-sensitive cultivar Fajr30. In other words, the tolerant cultivar produces more leaves but smaller under stress condition which finally reduces the leaf area compared to the sensitive cultivar. This can be a good solution for drought resistance through reduce evaporation from the leaf surface as well as shading. In addition, osmotic regulators such as proline and soluble sugars and the activity of antioxidant enzymes including catalase and peroxidase increased under drought stress and it was higher in tolerant cultivar. In fact, it seems that tolerant cultivar can tolerate drought stress through activating their immune system by producing osmoprotectant and increasing the activity of antioxidant enzymes.
Conclusion
In general, results showed that proline, soluble sugars and antioxidant enzymes play a role in the mechanism of stress tolerance and their metabolism is affected by drought stress. The results of this experiment suggest that the accumulation of these osmoprotectans and morphological changes can be part of the drought resistance mechanisms in the drought tolerant genotype of barley, which can ultimately be used in breeding programs to improve drought tolerance. In general, the results of this experiment showed that both barley cultivars responded to drought stress, but Dasht cultivar showed more tolerance in these conditions.

Keywords

Main Subjects

 
Abdel-Motagally, F.M.F., El-Zohri, M., 2018. Improvement of wheat yield grown under drought stress by boron foliar application at different growth stages. Journal of the Saudi Society of Agricultural Sciences. 17, 178-185. https://doi.org/10.1016/j.jssas.2016.03.005.
Abid, M., Tian, Z., Ata-Ul-Karim, S.T., Cui, Y., Liu, Y., Zahoor, R., Jiang, D., Dai, T., 2016. Nitrogen nutrition improves the potential of wheat (Triticum aestivum l.) to alleviate the eects of drought stress during vegetative growth periods. Frontiers in Plant Science 7, 981. https://doi.org/10.3389/fpls.2016.00981
Ahmadi, H., Abbasi, A., Taleei, A., Mohammadi, V., Pueyo, J.J., 2022. Antioxidant response and calcium-dependent protein kinases involvement in canola (Brassica napus L.) Tolerance to Drought. Agronomy. 12, 125.  https://doi.org/10.3390/agronomy12010125
Anjorin, F.B., Adejumo, S.A., Agboola, L., Samuel, Y.D., 2016. Proline, soluble sugar, leaf starch and relative water contents of four maize varieties in response to different watering regimes. Cercetari Agronomice in Moldova. 3, 51-62. https://doi.org/10.1515/cerce-2016-0025
Arendt, E.K., Zannini, E., 2013. Barley. In: Arendt, E.K., Zannini, E. (eds.), Cereal Grains for the Food and Beverage Industries. Woodhead Publishing, PP. 155-201.
Bakhshi Khaniki, G., Fatahi, F., Yazdchi, S., 2007. Drought effects of morphologic traits of 10 barley varieties in osko area, eastern azarbaijan province. Pajouhesh-Va-Sazandegi. 20, 108-114. [In Persian].
Bates, L.S., Waldren, R.P., Teare, I.D., 1973. Rapid determination of free proline for water stress studies. Plant and Soil. 39, 205-207. https://doi.org/10.1007/BF00018060
Bielach, A., Hrtyan, M., Tognetti, V.B., 2017. Plants under stress: involvement of auxin and cytokinin. International Journal of Molecular Sciences.18,1427. https://doi.org/10.3390/ijms18071427
Borzooei, A., Khazaie, H.R., Shariari, F., 2006. The impact of post anthesis drought stress on physiologycal characteristics and antioxidant enzymes in different wheat cultivars. Journal of Agricultural Science and Technology. 20, 65-75. [In Persian].
Brersen, C.R., Roddy, A.B., Wason, J.W., McElrone, A.J., 2019. Functional status of xylem through time. Annual Review of Plant Biology. 70, 407-433. https://doi.org/10.1146/annurev-arplant-050718-100455
Cai, K., Chen, X., Han, Z., Wu, X., Zhang, S., Li, Q., Zeng, F., 2020. Screening of worldwide barley collection for drought tolerance: the assessment of various physiological measures as the selection criteria. Frontiers in Plant Science. 11, 1159.  https://doi.org/10.3389/fpls.2020.01159
Chen, D., Wang, S., Cao, B., Cao, D., Leng, G., Li, H., Deng, X., 2016. Genotypic variation in growth and physiological response to drought stress and re-watering reveals the critical role of recovery in drought adaptation in maize seedlings. Frontiers in Plant Science 6, 1241. https://doi.org/10.3389/fpls.2015.01241
Darabi, M., Dashti, F., Gholami, M., Mosadeghi, M., Mirfattah. M., 2011. Effects of drought stress on yield and some morphological and physiological characteristics of tareh Irani (Allium ampeloperasum Tareh Group). Iranian Journal of Horticultural Science. 42, 95-103. [In Persian].
Dien, D. C., Mochizuki, T., Yamakawa, T., 2019. Effect of various drought stresses and subsequent recovery on proline, total soluble sugar and starch metabolisms in Rice (Oryza sativa L.) varieties. Plant Production Science. 22, 530-545. https://doi.org/10.1080/1343943X.2019.1647787
Dubois, D., Gilleres, K.A., Hamilton, J. K. 1956. Colorimetric method for determination of sugars and related substances. Analytical Chemistry. 28, 350-356. https://doi.org/10.1021/ac60111a017
FAO. 2021. FAOSTAT-Production/Crops. Retrieved March 3, 2021, from: http://www.fao.org/faostat/en/data/QC 
Farooq, M., Wahid, A., Kobayashi, N., Fujita, D., Basra, S., 2009. Plant drought stress: effects, mechanisms and management. In:  Lichtfouse, E., Navarrete, M., Debaeke, P., Veronique, S., Alberola, C. (eds.), Sustainable agriculture. Springer, Dordrecht, PP. 153-188
Ghotbi-Ravandi, A.A., Shahbazi, M., Shariati, M., Mulo, P., 2014. Effects of mild and severe drought stress on photosynthetic efficiency in tolerant and susceptible barley (Hordeum vulgare L.) genotypes. Journal of Agronomy and Crop Science. 200, 403-415. https://doi.org/10.1111/jac.12062
Gray, S.B., Brady, S.M., 2016. Plant developmental responses to climate change. Developmental biology. 419, 64-77. https://doi.org/10.1016/j.ydbio.2016.07.023.
Hafez, E.M., Seleiman, M.F., 2017. Response of barley quality traits, yield and antioxidant enzymes to water-stress and chemical inducers. International Journal of Plant Production. 1, 477-490. https://doi.org/10.22069/IJPP.2017.3712
Hafez, Y.M., Bacso, R., Kiraly, Z., Kunstler, A., Kiraly, L., 2012. Up-regulation of antioxidants in tobacco by low concentrations of H2O2 suppresses necrotic disease symptoms. Phytopathology Journal. 102, 848-856. https://doi.org/10.1094/PHYTO-01-12-0012-R
Harb, A., Simpson, C., Guo, W., Govindan, G., Kakani, V.G., Sunkar, R., 2020. The effect of drought on transcriptome and hormonal profiles in barley genotypes with contrasting drought tolerance. Frontiers in Plant Science. 11, 618491. https://doi.org/10.3389/fpls.2020.618491
Istanbuli, T., Baum, M., Touchan, H., Hamwieh, A., 2020. Evaluation of morpho physiological traits under drought stress conditions in barley (Hordeum vulgare L.). Photosynthetica. 58, 1059-1067. https://doi.org/10.32615/ps.2020.041
Jaleel, C.A., Manivannan, P., Wahid, A., Farooq, M., Al-Juburi, H.J., Somasundaram, R., Panneerselvam, R., 2009. Drought stress in plants: a review on morphological characteristics and pigments composition. International Journal of Agriculture and Biology. 11, 100-105.
Kamarudin, Z.S., Yusop, M.R., Tengku Muda Mohamed, M., Ismail, M.R., Harun, A.R., 2018. Growth performance and antioxidant enzyme activities of advanced mutant rice genotypes under drought stress condition. Agronomy. 8, 279. https://doi.org/10.3390/agronomy8120279
Kazerani, B., Navabpour, S., Sabouri, H., Ramezanpour, S.S., Zaynali Nezhad, K., Eskandari, A., 2019. Evaluation of proline content and enzymatic defense mechanism in response to drought stress in rice. Iranian journal of Plant Physiology. 9, 2749-2757. http://doi.org/10.30495/IJPP.2019.664580
Keyvan, S., 2010. The effects of drought stress on yield, relative water content, proline, soluble carbohydrates and chlorophyll of bread wheat cultivars. Journal of Animal and Plant Sciences. 8, 1051-1060.
Khan, M.N., Zhang, J., Luo, T., Liu, J., Ni, F., Rizwan, M., Hu, L., 2019. Morpho-physiological and biochemical responses of tolerant and sensitive rapeseed cultivars to drought stress during early seedling growth stage. Acta Physiologiae Plantarum. 41, 1-13. https://doi.org/10.1007/s11738-019-2812-2.
Khodabin, G., Tahmasebi‐Sarvestani, Z., Rad, A.H.S., Modarres‐Sanavy, S.A.M., 2020. Effect of drought stress on certain morphological and physiological characteristics of a resistant and a sensitive canola cultivar. Chemistry & Biodiversity. 17, e1900399.https://doi.org/10.1002/cbdv.201900399
Kumar Joshi, R., Nayak, S., 2010. Gene pyramiding-A broad spectrum technique for developing durable stress resistance in crops. Biotechnology and Molecular Biology Reviews. 5, 51–60.
Li, R.H., Guo, P.G., Michael, B., Stefania, G., Salvatore, C., 2006. Evaluation of chlorophyll content and fluorescence parameters as indicators of drought tolerance in barley. Agricultural Sciences in China. 5, 751-757. https://doi.org/10.1016/S1671-927(06)60120-X
Lichtenthaler, H.K., 1987. Chlorophylls and carotenoids: pigments of photosynthetic membranes. Methods in Enzymology. 148, 350–382. https://doi.org/10.1016/0076-6879(87)48036-1
Mejri, M., Siddique, K. H., Saif, T., Abdelly, C., Hessini, K., 2016. Comparative effect of drought duration on growth, photosynthesis, water relations, and solute accumulation in wild and cultivated barley species. Journal of Plant Nutrition and Soil Science. 179, 327-335. https://doi.org/10.1002/jpln.201500547
Movludi, A., Ebadi, A., Jahanbakhsh, S., Davari, M., Parmoon, G., 2014. The effect of water deficit and nitrogen on the antioxidant enzymes’ activity and quantum yield of barley (Hordeum vulgare L.). Notulae Botanicae Horti Agrobotanici Cluj-Napoca .42, 398-404. https://doi.org/10.15835/nbha4229340
Naeemi, T., Fahmideh, L., Fakheri, B., 2018. The impact of drought stress on antioxidant enzymes activities, containing of proline and carbohydrate in some genotypes of durum wheat (Triticum turgidu l.) at seedling stage. Journal of Crop Breeding. 10, 22-31. [In Persian].
Noctor, G., Mhamdi, A., Foyer, C., 2014. Roles of reactive oxygen metabolism in drought: not so cut and dried. Plant Physiology. 164, 1636-1648. https://doi.org/10.1104/pp.113.233478
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. https://doi.org/10.3923/ajpp.2010.338.349
Qaseem, M. F., Qureshi, R., Shaheen, H., 2019. Effects of pre-anthesis drought, heat and their combination on the growth, yield and physiology of diverse wheat (Triticum aestivum L.) genotypes varying in sensitivity to heat and drought stress. Scientific Reports. 9, 1-12. https://doi.org/10.1038/s41598-019-43477-z
Rouhani, L., Zamani, M.J., Fotovat, R., 2015. Variation in stomatal size and density of barley genotypes under drought stress and normal conditions. Journal of Plant Research (Iranian Journal of Biology) 28, 986-994. [In Persian].
Sallam, A., Alqudah, A.M., Dawood, M.F., Baenziger, P.S., Borner, A., 2019. Drought stress tolerance in wheat and barley: advances in physiology, breeding and genetics research. International Journal of Molecular Sciences. 20, 31-37. https://doi.org/10.3390/ijms20133137
Samarah, N.H., Alqudah, A.M., Amayreh, J.A., McAndrews, G.M., 2009. The effect of late‐terminal drought stress on yield components of four barley cultivars. Journal of Agronomy and Crop Science. 195, 427-441. https://doi.org/10.1111/j.1439-037X.2009.00387.x
Seed and Plant Research Improvement Institute., 2015. Introduction of crop cultivars. Agricultural Research Education and Extention Organization 1-231. [In Persian].
Shabani, Z., Akbari, G.H., KhoshKholgh Sima., N.A., 2012. Effect of terminal drought stress on proline and soluble carbohydrate accumulation in sensitive and tolerant barley genotypes.p. 1-12. Proceedings of the 12th Iranian Crop Sciences Congress.  4 Sep. 2012. Karaj, Iran.
Sikuku, P.A., Netondo G.W., Onyango J.C., Musyimi D.M., 2010. Chlorophyll fluorescence, protein and chlorophyll content of three nerica rainfed rice varieties under varying irrigation regimes. ARPN Journal of Agricultural and Biological Science. 5, 19-25.
Smart, R.E., Bingham G.E., 1974. Rapid estimates of relative water content. Plant Physiology. 53, 258–260. https://doi.org/10.1104/pp.53.2.258
Srinivas, N.D., Rashmi K.R., Raghavarao K.S.M.S., 1999. Extraction and purification of a plant peroxidase by aqueous two-phase extraction coupled with gel filtration. Process Biochemistry. 35, 43–48. https://doi.org/10.1016/S0032-9592(99)00030-8
Velikova, V., Yordanov, I., Edreva, A., 2000. Oxidative stress and some antioxidant systems in acid rain-treated bean plants: protective roles of exogenous polyamines. Plant Science. 151, 59-66. https://doi.org/10.1016/S0168-9452(99)00197-1
Vinocur, B., Altman, A., 2005. Recent advances in engineering plant tolerance to abiotic stress: Achievements and limitations. Current Opinion in Biotechnology .16 12,3–132. https://doi.org/10.1016/j.copbio.2005.02.001
Yang, S., Deng, X., 2015. Effects of drought stress on antioxidant enzymes in seedlings of different wheat genotypes. Pakistan Journal of Botany. 47, 49-56.
Ying, Y.Q., Song, L.L., Jacobs, D.F., Mei, L., Liu, P., Jin, S.H., Wu, J.S., 2015. Physiological response to drought stress in Camptotheca acuminata seedlings from two provenances. Frontiers in Plant Science. 6, 361. https://doi.org/10.3389/fpls.2015.00361
Zahedi, M. B., Razi, H., Saed-Moucheshi, A., 2016. Evaluation of antioxidant enzymes, lipid peroxidation and proline content as selection criteria for grain yield under water deficit stress in barley. Journal of Applied Biological Sciences. 10, 71-78.