Environmental Stresses in Crop Sciences

Environmental Stresses in Crop Sciences

Evaluation of the response of sesame (Sesamum indicum L.) genotypes to drought stress

Articles in Press

Document Type : Original Article

Authors
Sakineh Padiab 1
Rasool Asghari Zakaria 2
Nasser Zare 2
Nooraddin Hosseinpour َAzad 3
Mohammad Zaeifizadeh 4
Parisa Sheikhzadeh 5
1 Ph.D. Student of Agricultural Biotechnology, Department of Plant Production and Genetics, Faculty of Agriculture and Natural Resources, University of Mohaghegh Ardabili, Ardabil, Iran
2 Professor, Department of Plant Production and Genetics, Faculty of Agriculture and Natural Resources, University of Mohaghegh Ardabili, Ardabil, Iran
3 Associate Professor, Department of Plant Sciences and Medicinal Plants, Faculty of Agriculture, Meshginshahr, University of Mohaghegh Ardabili, Ardabil, Iran
4 Associate Professor, Department of Biology, Ardabil Branch, Islamic Azad University, Ardabil, Iran
5 Associate Professor, Department of Plant Production and Genetics, Faculty of Agriculture and Natural Resources, University of Mohaghegh Ardabili, Ardabil, Iran
10.22077/escs.2026.9381.2368
Abstract
Introduction
Sesame (Sesamum indicum L.) is one of the world's most important oilseed crops and is highly susceptible to abiotic stresses, particularly drought. Drought reduces plant growth and yield and alters morphological and biochemical characteristics. This study aimed to evaluate the responses of different sesame genotypes to drought stress and identify drought-tolerant genotypes suitable for cultivation in arid regions.
 
Materials and methods
In 2023, an experiment was conducted in the growth chamber of the Faculty of Agriculture at Mohaghegh Ardabili University to evaluate the responses of 10 sesame genotypes (Halil, Dashtestan, Darab, Sardari, Line 24, Line 25, Line 26, Oltan, Naz-Tekshakheh, and Yellow White) to drought stress. The experiment was conducted as a factorial arrangement in a completely randomized design (CRD) with three replications under hydroponic conditions using perlite and Hoagland nutrient solution. Drought stress was imposed at the four-leaf stage using polyethylene glycol (PEG 6000) at three levels (0, −3, and −6 bar). Shoot samples were collected to assess morphological traits, biochemical parameters (proline, soluble carbohydrates, hydrogen peroxide, malondialdehyde, total phenolic content, and total flavonoid content), and polyphenol oxidase activity. Data were analyzed using STAR software, and treatment means were compared using the least significant difference (LSD) test at the 1% probability level. The yield stability index (YSI) and membership function value (MFV) were used to evaluate genotype stability under drought stress.
 
Results and discussion
Evaluation of sesame genotypes under drought stress showed that drought significantly reduced morphological traits, including seedling length and dry weight, in all genotypes. However, the Sardari genotype exhibited the smallest reductions and performed best under both drought stress levels. In contrast, proline and soluble carbohydrate contents increased with increasing drought intensity. The greatest proline accumulation was observed in the Yellow White and Darab genotypes, whereas Sardari exhibited the highest soluble carbohydrate content. Oxidative stress markers, including hydrogen peroxide (H₂O₂) and malondialdehyde (MDA), also increased under drought stress. Dashtestan and Line 25 exhibited the highest levels of oxidative stress markers, indicating greater oxidative damage, whereas Oltan and Darab had the lowest levels. Total phenolic and total flavonoid contents reached their highest levels under moderate drought stress. Yellow White, Darab, Sardari, and Line 24 exhibited greater accumulation of these metabolites, whereas Halil and Naz-Tekshakheh showed lower levels. Among the antioxidant enzymes, catalase activity was highest in Line 25, Line 26, and Sardari under drought stress. Moreover, Naz-Tekshakheh and Sardari exhibited the highest polyphenol oxidase activity under severe drought stress, whereas Yellow White and Halil exhibited the lowest activity. Analysis of YSI and MFV indicated that Sardari and Darab had the highest YSI and MFV values under both drought stress levels and were identified as the most drought-tolerant genotypes. In contrast, Line 25 and Halil performed poorly across most evaluated traits and were classified as drought-sensitive genotypes.
 
Conclusion
The findings demonstrated that drought stress significantly affected the morphological and biochemical characteristics of sesame genotypes, although the extent of these effects varied among genotypes. Sardari and Darab consistently exhibited superior performance under both moderate and severe drought stress, as evidenced by smaller reductions in morphological traits, greater accumulation of osmoprotectants (proline and soluble carbohydrates), lower levels of oxidative stress markers, and stronger antioxidant enzyme activities. These genotypes also had the highest values for the drought tolerance indices (YSI and MFV), indicating greater stability and adaptability under water-limited conditions. Conversely, Line 25 and Halil were more susceptible to drought stress, exhibiting greater growth inhibition and increased oxidative damage. Overall, Sardari and Darab can be recommended as promising candidates for cultivation and breeding programs aimed at enhancing drought tolerance in sesame, especially in arid and semi-arid regions.
Keywords
Defense mechanisms
Osmotic stress
Polyethylene glycol
Reactive Oxygen species
Subjects
Afzal, S., Chaudhary, N., Singh, N.K., 2021. Role of soluble sugars in metabolism and sensing under abiotic stress. In: Aftab, T., Hakeem, K.R. (Eds.), Plant Growth Regulators. Springer, Cham, Switzerland. https://doi.org/10.1007/978-3-030-61153-8_14
Ahmad, F., Singh, A., Kamal, A., 2020. Osmoprotective role of sugar in mitigating abiotic stress in plants. In: Roychoudhury, A., Tripathi, D.K. (eds.), Protective Chemical Agents in the Amelioration of Plant Abiotic Stress: Biochemical and Molecular Perspectives. Wiley Online Library, pp. 53–70. https://doi.org/10.1002/9781119552154.ch3
Ahmed, M., Kheir, A.M., Mehmood, M.Z., Ahmad, S., Hasanuzzaman, M., 2022. Changes in germination and seedling traits of sesame under simulated drought. Phyton. 91, 713–726. https://doi.org/10.32604/phyton.2022.018552
Apel, K., Hirt, H., 2004. Reactive oxygen species: Metabolism, oxidative stress, and signal transduction. Annual Review of Plant Biology. 55, 373–399.    https://doi.org/10.1146/annurev.arplant.55.031903.141701
Arora, M., Saxena, P., Abdin, M.Z., Varma, A., 2020. Interaction between Piriformospora indica and Azotobacter chroococcum diminishes the effect of salt stress in Artemisia annua L. by enhancing enzymatic and non-enzymatic antioxidants. Symbiosis. 80, 61–73. https://doi.org/10.1007/s13199-019-00656-w
Badea, A., Tucker, J.R., Sabra, A., Netticadan, T., Blackwell, B., Yu, L., O'Sullivan, D.M., 2023. Endogenic phenolic compounds of barley as potential biomarkers related to grain mycotoxin production and cultivar selection. Biology. 12, 1306. https://doi.org/10.3390/biology12101306
Bahrami, H., Razmjoo, J., Jafari, A.O., 2012. Effect of drought stress on germination and seedling growth of sesame cultivars (Sesamum indicum L.). International Journal of AgriScience. 2, 423–428.
Bate, N.J., Roth, S.A., 1973. The determination of proline in plant tissues. Plant Physiology. 51, 917–921. https://doi.org/10.1104/pp.51.5.917
Bouslama, M., Schapaugh, W.T., 1984. Stress tolerance in soybean. Part 1: Evaluation of three screening techniques for heat and drought tolerance. Crop Science. 24, 933–937. https://doi.org/10.2135/cropsci1984.0011183X002400050026x
Chakraborty, K., Bishi, S.K., Goswami, N., Singh, A.L., Zala, P.V., 2016. Differential fine-regulation of enzyme driven ROS detoxification network imparts salt tolerance in contrasting peanut genotypes. Environmental and Experimental Botany. 128, 79-90. https://doi.org/10.1016/j.envexpbot.2016.05.001
Chang, C., Yang, M., Wen, H., Chern, J., 2002. Estimation of total flavonoid content in propolis by two complementary colorimetric methods. Journal of Food and Drug Analysis. 10, 178–182. https://doi.org/10.38212/2224-6614.2748
Chen, D., Mubeen, B., Hasnain, A., Rizwan, M., Adrees, M., Naqvi, S.H., Batool, M., Kamran, M., El-Sappah, A.H., 2022. Role of promising secondary metabolites to confer resistance against environmental stresses in crop plants: Current scenario and future perspectives. Frontiers in Plant Science. 13, 881032. https://doi.org/10.3389/fpls.2022.881032
Chen, W., Zhang, H., Zhang, G., Wu, F., Zhou, M., 2012. Screening of barley genotypes for salt tolerance based on germination, seedling growth and physiological responses. Soil Science and Plant Nutrition. 58, 219–230. https://doi.org/10.1080/00380768.2012.669002
Dehghanian, Z., Habibi, K., Dehghanian, M., Aliyar, S., Lajayer, B. A., Astatkie, T., Minkina, T., Keswani, C., 2022. Reinforcing the bulwark: Unravelling the efficient applications of plant phenolics and tannins against environmental stresses. Heliyon. 8, e09094. https://doi.org/10.1016/j.heliyon.2022.e09094
Dossa, K., Niang, M., Assogbadjo, A.E., Cissé, N., Diouf, D., 2016. Whole genome homology-based identification of candidate genes for drought resistance in Sesamum indicum L. African Journal of Biotechnology. 15, 1464–1475. https://doi.org/10.5897/AJB2016.15420
Dossa, K., Yehouessi, L. W., Likeng-Li-Ngue, B. C., Diouf, D., Liao, B., Zhang, X., Cissé, N., Bell, J.M., 2017. Comprehensive screening of some West and Central African sesame genotypes for drought resistance: probing by agromorphological, physiological, Biochemical, and seed quality traits. Agronomy. 7, 83. https://doi.org/10.3390/agronomy7040083
Dreccer, M.F., Fainges, J., Whish, J., Ogbonnaya, F.C., Sadras, V.O., 2018. Comparison of sensitive stages of wheat, barley, canola, chickpea, and field pea to temperature and water stress across Australia. Agricultural and Forest Meteorology. 248, 275–294.      https://doi.org/10.1016/j.agrformet.2017.10.006
Du, Y., Zhao, Q., Chen, L., Yao, X., Zhang, H., Wu, J., Xie, F., 2020. Effect of drought stress during soybean R2–R6 growth stages on sucrose metabolism in leaf and seed. International Journal of Molecular Sciences. 21, 618. https://doi.org/10.3390/ijms21020618
Dubois, M., Gilles, K.A., Hamilton, J.K., Rebers, P.A., Smith, F., 1956. Colorimetric method for determination of sugars and related substances. Analytical Chemistry. 28, 350–356. https://doi.org/10.1021/ac60111a017
Ebrahimian, E., Seyyedi, S.M., Bybordi, A., Damalas, C.A., 2019. Seed yield and oil quality of sunflower, safflower, and sesame under different levels of irrigation water availability. Agricultural Water Management. 218, 149–157. https://doi.org/10.1016/j.agwat.2019.03.031
Eskandari, H., Zehtab-Salmasi, S., Ghassemi-Golezani, K., Gharineh, M.H., 2009. Effects of water limitation on grain and oil yields of sesame cultivars. Journal of Food, Agriculture & Environment 7, 339–342.
Fraire-Velázquez, S., Balderas-Hernández, V. E., 2013. Abiotic stress in plants and metabolic responses. In: Vahdati, K., Leslie, C. (Eds.), Abiotic Stress – Plant Responses and Applications in Agriculture. InTech Open. https://doi.org/10.5772/54859
Gholamhoseini, M., 2020. Evaluation of sesame genotypes for agronomic traits and stress indices grown under different irrigation treatments. Agronomy Journal. 112, 1794–1804. https://doi.org/10.1002/agj2.20167
Hanif, S., Saleem, M.F., Sarwar, M., Irshad, M., Shakoor, A., Wahid, M.A., Khan, M.A., 2021. Biochemically triggered heat and drought stress tolerance in rice by proline application. Journal of Plant Growth Regulation. 40, 305–312. https://doi.org/10.1007/s00344-020-10095-3
Hasanuzzaman, M., Hossain, M.A., Teixeira da Silva, J.A., Fujita, M., 2012. Plant responses and tolerance to abiotic oxidative stress: Antioxidant defense is a key factor. In: Bandi, V., Shanker, A.K., Shanker, C., Mandapaka, M. (eds.), Crop Stress and Its Management: Perspectives and Strategies. Springer, pp. 261–316.
Hasanuzzaman, M., Nahar, K., Anee, T.I., Khan, M.I.R., Fujita, M., 2018. Silicon-mediated regulation of antioxidant defense and glyoxalase systems confers drought stress tolerance in Brassica napus L. South African Journal of Botany. 115, 50–57.           https://doi.org/10.1016/j.sajb.2017.12.006
Hatzig, S.V., Nuppenau, J.N., Snowdon, R.J., Schießl, S.V., 2018. Drought stress has transgenerational effects on seeds and seedlings in winter oilseed rape (Brassica napus L.). BMC Plant Biology. 18, 297. https://doi.org/10.1186/s12870-018-1531-y
Heath, R. L., Packer, L., 1968. Photoperoxidation in isolated chloroplasts. I. Kinetics and stoichiometry of fatty acid peroxidation. Archives of Biochemistry and Biophysics. 125, 189–198. https://doi.org/10.1016/0003-9861(68)90028-4
Hellal, F.A., El-Shabrawi, H.M., El-Hady, A.M., Khatab, I.A., El-Sayed, S.A.A., Abdelly, C., 2018. Influence of PEG-induced drought stress on molecular and biochemical constituents and seedling growth of Egyptian barley cultivars. Journal of Genetic Engineering and Biotechnology. 16, 203–212. https://doi.org/10.1016/j.jgeb.2017.10.009
Hoyos, B.E., Johnson, J.B., Mani, J.S., Batley, R.J., Trotter, T., Bhattarai, S.P., Naiker, M., 2024. The effect of water stress on bioactive compounds in Australian-grown black sesame. Plants. 13, 793. https://doi.org/10.3390/plants13060793
Hussain, H.A., Men, S., Hussain, S., Chen, Y., Ali, S., Zhang, S., Zhang, K., Li, Y., Xu, Q., Liao, C., Wang, L., 2019. Interactive effects of drought and heat stresses on morpho-physiological attributes, yield, nutrient uptake and oxidative status in maize hybrids. Scientific Reports. 9, 3890. https://doi.org/10.1038/s41598-019-40362-7
Islam, F., Yasmeen, T., Ali, Q., Ali, S., Arif, M. S., Hussain, S., Rizvi, H., 2014. Influence of Pseudomonas aeruginosa as PGPR on oxidative stress tolerance in wheat under Zn stress. Ecotoxicology and Environmental Safety. 104, 285–293.          https://doi.org/10.1016/j.ecoenv.2014.03.008
Jatav, S.K., Kandalkar, V.S., 2014. Assessment of wheat genotypes for yield potential and stress adaptation. Journal of Wheat Research. 6, 29–36.            http://epubs.icar.org.in/ejournal/index.php/JWR
Kabbadj, A., Makoudi, B., Mouradi, M., Pauly, N., Frendo, P., Ghoulam, C., 2017. Physiological and biochemical responses involved in water deficit tolerance of nitrogen-fixing Vicia faba. PLoS ONE. 12, e0190284. https://doi.org/10.1371/journal.pone.0190284
Kadkhodaie, A., Razmjoo, J., Zahedi, M., Pessarakli, M., 2014. Oil content and composition of sesame (Sesamum indicum L.) genotypes as affected by irrigation regimes. Journal of the American Oil Chemists’ Society. 91, 1737–1744. https://doi.org/10.1007/s11746-014-2524-0
Kaushal, M., Wani, S.P., 2016. Rhizobacterial-plant interactions: Strategies ensuring plant growth promotion under drought and salinity stress. Agriculture, Ecosystems and Environment. 231, 68–78. https://doi.org/10.1016/j.agee.2016.06.031
Kaya, M.D., Okçu, G., Atak, M., Çıkılı, Y., Kolsarıcı, Ö., 2006. Seed treatments to overcome salt and drought stress during germination in sunflower (Helianthus annuus L.). European Journal of Agronomy. 24, 291–295. https://doi.org/10.1016/j.eja.2005.08.001
Kazemi Oskuei, B., Bandehagh, A., Farajzadeh, D., Asgari Lajayer, B., Rajput, V.D., Astatkie, T., 2023. Morphological, biochemical, and physiological responses of canola cultivars to drought stress. International Journal of Environmental Science and Technology. 20, 13551–13560. https://doi.org/10.1007/s13762-023-04928-3
Khan, P., Abdelbacki, A.M.M., Albaqami, M., Jan, R., Kim, K.-M., 2025. Proline promotes drought tolerance in maize. Biology. 14, 41. https://doi.org/10.3390/biology14010041
Khanna-Chopra, R. Selote, D.S., 2007. Acclimation to drought stress generates oxidative stress tolerance in drought-resistant than -susceptible wheat cultivars under field conditions. Environmental and Experimental Botany. 60, 276–283. https://doi.org/10.1016/j.envexpbot.2006.11.004
Khazaei, Z., Estaji, A., 2020. Effect of foliar application of ascorbic acid on sweet pepper (Capsicum annuum) plants under drought stress. Acta Physiologiae Plantarum. 42, 1–12. https://doi.org/10.1007/s11738-020-03106-z
Kouighat, M., Moussaoui, F.E., Adiba, A., Hafid, A., Bouchyoua, A., Fechtali, M.E., Nabloussi, A., 2025. Evaluation and selection of sesame mutants (Sesamum indicum L.) for optimal nutritional profiles in seeds under field drought conditions. Oilseeds and Fats Crops and Lipid-OCL. 32. https://doi.org/10.1051/ocl/2025004
Ku, Y.S., Au-Yeung, W.K., Yung, Y.L., Li, M.W., Wen, C.Q., Liu, X., Lam, H.M., 2013. Drought stress and tolerance in soybean. In: Board, J. (ed.), A Comprehensive Survey of International Soybean Research: Genetics, Physiology, Agronomy and Nitrogen Relationships. InTech. pp. 209–237. https://doi.org/10.5772/45867   
Liu, C., Yang, Z., Hu, Y.G., 2015. Drought resistance of wheat alien chromosome addition lines evaluated by membership function value based on multiple traits and drought resistance index of grain yield. Field Crops Research. 179, 103–112.
Liu, Y.N., Xu, Q.Z., Li, W.C., Yang, X.H., Zheng, Q., Li, B., Li, Z.S., Li, H.W., 2019. Long-term high light stress induces leaf senescence in wheat (Triticum aestivum L.). Photosynthetica. 57, 830–840. https://doi.org/10.32615/ps.2019.086
Mahdavi Khorami, A., Masoud Sinaki, J., Amini Dehaghi, M., Rezvan, S., Damavandi, A., 2019. Sesame (Sesamum indicum L.) biochemical and physiological responses as affected by applying chemical, biological, and nano-fertilizers in field water stress conditions. Journal of Plant Nutrition. 43, 456–475. https://doi.org/10.1080/01904167.2019.1683189
Mahpara, S., Zainab, A., Ullah, R., Kausar, S., Bilal, M., Latif, M.I., Zuan, A.T.K., 2022. The impact of PEG-induced drought stress on seed germination and seedling growth of different bread wheat (Triticum aestivum L.) genotypes. PLoS ONE. 17, e0262937.         https://doi.org/10.1371/journal.pone.0262937
Manghwar, H., Hussain, A., Alam, I., Khoso, M. A., Ali, Q., Liu, F., 2024. Waterlogging stress in plants: Unraveling the mechanisms and impacts on growth, development, and productivity. Environmental and Experimental Botany. 105824.       https://doi.org/10.1016/j.envexpbot.2024.105824
Marinova, D., Ribarova, F., Atanassova, M., 2005. Total phenolics and total flavonoids in Bulgarian fruits and vegetables. Journal of the University of Chemical Technology and Metallurgy. 40, 255–260. https://doi.org/10.13140/2.1.1413.1841
Marmar, A., Baenziger, S., Dweikat, I., El Hussein, A. A., 2013. Preliminary screening for water stress tolerance and genetic diversity in wheat (Triticum aestivum L.) cultivars from Sudan. Journal of Genetic Engineering and Biotechnology. 11, 87–94.  https://doi.org/10.1016/j.jgeb.2013.08.004
Martínez-Vilalta, J., Sala, A., Asensio, D., Galiano, L., Hoch, G., Palacio, S., Piper, F.I., Lloret, F., 2016. Dynamics of non-structural carbohydrates in terrestrial plants: A global synthesis. Ecological Monographs. 86, 495–516. https://doi.org/10.1002/ecm.1231
Mi, N., Cai, F., Zhang, Y., Ji, R., Zhang, S., Wang, Y., 2018. Differential responses of maize yield to drought at vegetative and reproductive stages. Plant, Soil and Environment. 64, 260–267. https://doi.org/10.17221/141/2018-PSE
Michel, B.E., Kaufmann, M.R., 1973. The osmotic potential of polyethyleneglycol 6000. Plant Physiology. 51, 914–916. https://doi.org/10.1104/pp.51.5.914
Mohan, N., Kumari, N., Jattan, M., Avtar, R., Rathore, V., 2020. Aftermath of terminal heat stress on Indian mustard (Brassica juncea L.): A brief review. Journal of Oilseed Brassica, 11, 1–8.
Mohi-Ud-Din, M., Hossain, M., Rohman, M., Uddin, M., Haque, M., Ahmed, J.U., Mostofa, M.G., 2021. Multivariate analysis of morpho-physiological traits reveals differential drought tolerance potential of bread wheat genotypes at the seedling stage. Plants. 10, 879.        https://doi.org/10.3390/plants10050879
Muhammad, M., Waheed, A., Wahab, A., Majeed, M., Nazim, M., Liu, Y.-H., Li, L., Li, W.-J., 2023. Soil salinity and drought tolerance: An evaluation of plant growth, productivity, microbial diversity, and amelioration strategies. Plant Stress. 11, 100319.          https://doi.org/10.1016/j.stress.2023.100319
Mwadzingeni, L., Shimelis, H., Tesfay, S., Tsilo, T. J., 2016. Screening of bread wheat genotypes for drought tolerance using phenotypic and proline analyses. Frontiers in Plant Science. 7, 1276. https://doi.org/10.3389/fpls.2016.01276
Ngompe Deffo, T., Kouam, E.B., Mandou, M.S., Bara, R.A., Chotangui, A.H., Souleymanou, A., Beyegue Djonko, H., Tankou, C.M., 2024. Identifying critical growth stage and resilient genotypes in cowpea under drought stress contributes to enhancing crop tolerance for improvement and adaptation in Cameroon. PLoS ONE. 19, e0304674. https://doi.org/10.1371/journal.pone.0304674
Ozkan, A., Kulak, M., 2013. Effects of water stress on growth, oil yield, fatty acid composition and mineral content of Sesamum indicum. Journal of Animal and Plant Sciences. 23, 1686–1690.
Ozkur, O., Ozdemir, F., Bor, M., Turkan, I., 2009. Physiochemical and antioxidant responses of the perennial xerophyte Capparis ovata Desf. to drought. Environmental and Experimental Botany. 66, 487–492. https://doi.org/10.1016/j.envexpbot.2009.04.003
Pandey, P., Ramegowda, V., Senthil-Kumar, M., 2015. Shared and unique responses of plants to multiple individual stresses and stress combinations: Physiological and molecular mechanisms. Frontiers in Plant Science. 6, 723. https://doi.org/10.3389/fpls.2015.00723
Pathak, N., Rai, A.K., Kumari, R., Thapa, A., Bhat, K.V., 2014. Sesame crop: An underexploited oilseed holds tremendous potential for enhanced food value. Agricultural Sciences. 5, 519–529. https://doi.org/10.4236/as.2014.56054
Qureshi, M., Arslan, M., Golukcu, M., Bera, S. K., Uzun, B., Yol, E., 2023. Assessment of drought tolerance of sesame germplasm with agronomic and quality traits. Crop Science, 63, 2763–2777. https://doi.org/10.1002/csc2.21047
Rai, K.K., Pandey, N., Rai, S.P., 2020. Salicylic acid and nitric oxide signaling in plant heat stress. Physiologia Plantarum. 168, 241–255. https://doi.org/10.1111/ppl.12958
Rajametov, S.N., Yang, E.Y., Cho, M.C., Chae, S.Y., Jeong, H.B., Choi, K.Y., Lee, S.Y., Kim, Y.J., 2021. Heat tolerant hot pepper exhibits constant photosynthesis via increased transpiration rate, high proline content and fast recovery in heat stress condition. Scientific Reports. 11, 1–9. https://doi.org/10.1038/s41598-021-93697-5
Raymond, J., Rakariyatham, N., Azanza, J. L., 1993. Purification and some properties of polyphenol oxidase from sunflower seeds. Phytochemistry. 34, 927–931.          https://doi.org/10.1016/0031-9422(93)80039-T
Raza, M.A.S., Shahid, A.M., Saleem, M.F., Khan, I.H., Ahmad, S., Ali, M., Iqbal, R., 2017. Effects and management strategies to mitigate drought stress in oilseed rape (Brassica napus L.): a review. Zemdirbyste-Agriculture. 104, 85–94. https://doi.org/10.13080/z-a.2017.104.012
Reddy, A.R., Chaitanya, K.V., Vivekanandan, M., 2004. Drought-induced responses of photosynthesis and antioxidant metabolism in higher plants. Journal of Plant Physiology. 161, 1189–1202. https://doi.org/10.1016/j.jplph.2004.01.013
ReportLinker., 2024. Global Sesame Oil Market Size, Share and Trends Analysis Report. Retrieved from https://www.reportlinker.com/p06487366/Global-Sesame-Oil-Market-Size-Share-Trends-Analysis-Report.html
Rezayian, M., Ebrahimzadeh, H., Niknam, V., 2020. Nitric oxide stimulates antioxidant system and osmotic adjustment in soybean under drought stress. Journal of Soil Science and Plant Nutrition. 16, 1–11. https://doi.org/10.1007/s42729-020-00198-x
Seleiman, M.F., Al-Suhaibani, N., Ali, N., Akmal, M., Alotaibi, M., Refay, Y., Dindaroglu, T., Abdul-Wajid, H.H., Battaglia, M.L., 2021. Drought stress impacts on plants and different approaches to alleviate its adverse effects. Plants (Basel).10, 259.         https://doi.org/10.3390/plants10020259
Shahzad, A., Gul, H., Ahsan, M., Ahmad, M., Khan, S.A., Iqbal, M., 2023. Comparative genetic evaluation of maize inbred lines at seedling and maturity stages under drought stress. Journal of Plant Growth Regulation. 42, 989–1005. https://doi.org/10.1007/s00344-022-10608-2
Singh, S., Angadi, S.V., Grover, K., Begna, S., Auld, D., 2016. Drought response and yield formation of spring safflower under different water regimes in the semiarid southern high plains. Agricultural Water Management. 163, 354–362. https://doi.org/10.1016/j.agwat.2015.10.010
Singh, V.K., Singh, A.K., Singh, P.P., Kumar, A., 2018. Interaction of plant growth promoting bacteria with tomato under abiotic stress: A review. Agriculture, Ecosystems and Environment. 267, 129–140. https://doi.org/10.1016/j.agee.2018.08.020
Singhal, R.K., Jatav, H.S., Aftab, T., Pandey, S., Mishra, U.N., Chauhan, J., Khan, M.I.R., 2021. Roles of nitric oxide in conferring multiple abiotic stress tolerance in plants and crosstalk with other plant growth regulators. Journal of Plant Growth Regulation. 40, 2303–2328. https://doi.org/10.1007/s00344-021-10446-8
Song, R., Shi, P., Xiang, L., He, Y., Dong, Y., Miao, Y., Qi, J., 2024. Evaluation of barley genotypes for drought adaptability: based on stress indices and comprehensive evaluation as criteria. Frontiers in Plant Science. 15, 1436872. https://doi.org/10.3389/fpls.2024.1436872
Sun, J., Rao, Y., Le, Mei, Yan, T., Yan, X., Zhou, H., 2010. Effects of drought stress on sesame growth and yield characteristics and comprehensive evaluation of drought tolerance. Chinese Journal of Oil Crop Sciences. 32, 525–533.
Sun, Y., Robert, C.A.M., Thakur, M.P., 2024. Drought intensity and duration effects on morphological root traits vary across trait type and plant functional groups: A meta-analysis. BMC Ecology and Evolution. 24, 92. https://doi.org/10.1186/s12862-024-02275-6
Velikova, V., Yordanov, I., Edreva, A., 2000. Oxidative stress and some antioxidant systems in acid rain-treated bean plants: Protective role of exogenous polyamines. Plant Science. 151, 59–66. https://doi.org/10.1016/S0168-9452(99)00197-1
Were, B.A., Onkware, O.A., Gudu, S., Welander, M., Carlsson, A.S., 2006. Seed oil content and fatty acid composition in East African sesame (Sesamum indicum L.) accessions evaluated over 3 years. Field Crops Research. 97, 254–260. https://doi.org/10.1016/j.fcr.2005.10.009
Wiley, E., King, C.M., Landhäusser, S. M., 2019. Identifying the relevant carbohydrate storage pools available for remobilization in aspen roots. Tree Physiology. 39, 1109–1120. https://doi.org/10.1093/treephys/tpz051
Yang, X., Wang, B., Chen, L., Li, P., Cao, C., 2019. The different influences of drought stress at the flowering stage on rice physiological traits, grain yield, and quality. Scientific Reports. 9, 3742. https://doi.org/10.1038/s41598-019-40161-0
Zhang, J., Kirkham, M.B., 1996. Antioxidant responses to drought in sunflower and sorghum seedlings. New Phytologist. 132, 361–373. https://doi.org/10.1111/j.1469-8137.1996.tb01856.x
Zhang, Y., Xu, J., Li, R., Ge, Y., Li, Y., Li, R., 2023. Plants’ response to abiotic stress: Mechanisms and strategies. International Journal of Molecular Sciences. 24, 10915. https://doi.org/10.3390/ijms241310915
Zhu, X., Liu, W., Wang, B., Yang, L., 2024. Molecular and physiological responses of two quinoa genotypes to drought stress. Frontiers in Genetics. 15, 1439046. https://doi.org/10.3389/fgene.2024.1439046
Environmental Stresses in Crop Sciences

Articles in Press, Accepted Manuscript
Available Online from 25 July 2026

  • Receive Date 19 May 2025
  • Revise Date 23 June 2025
  • Accept Date 29 June 2025