{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,4]],"date-time":"2026-09-04T05:17:00Z","timestamp":1788499020831,"version":"build-2803163510"},"reference-count":153,"publisher":"MDPI AG","issue":"17","license":[{"start":{"date-parts":[[2026,8,29]],"date-time":"2026-08-29T00:00:00Z","timestamp":1787961600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["31701466"],"award-info":[{"award-number":["31701466"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"award":["31701466"],"award-info":[{"award-number":["31701466"]}],"id":[{"id":"https:\/\/ror.org\/01h0zpd94","id-type":"ROR","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100010650","name":"Huanggang Normal University","doi-asserted-by":"publisher","award":["2042023059"],"award-info":[{"award-number":["2042023059"]}],"id":[{"id":"10.13039\/501100010650","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100010650","name":"Huanggang Normal University","doi-asserted-by":"publisher","award":["204202505004"],"award-info":[{"award-number":["204202505004"]}],"id":[{"id":"10.13039\/501100010650","id-type":"DOI","asserted-by":"publisher"}]},{"award":["2042023059"],"award-info":[{"award-number":["2042023059"]}],"id":[{"id":"https:\/\/ror.org\/007gf6e19","id-type":"ROR","asserted-by":"publisher"}]},{"award":["204202505004"],"award-info":[{"award-number":["204202505004"]}],"id":[{"id":"https:\/\/ror.org\/007gf6e19","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>Wheat (Triticum aestivum L.) is considered one of the most important cereals globally, contributing significantly to the human population\u2019s caloric and protein requirements. Therefore, ensuring a sufficient yield of wheat for global consumption plays a significant role in maintaining food security in different parts of the world. With increasing demand and dwindling production capacity, due to increasingly uncertain growing conditions, projections indicate that there should be an upswing of 60\u201370% in wheat productivity by 2050 to fulfill the requirement. However, drought represents the most significant and widespread abiotic limitation to global wheat production, currently resulting in approximately 10% yield losses worldwide. Furthermore, each additional 1 \u00b0C increase in temperature is anticipated to decrease staple calorie production by 4.4%. The factors contributing to drought in wheat, as well as its impact on the plant\u2019s biochemical, physiological, and morphological structures, include altered rainfall patterns, elevated atmospheric CO2 levels, increased temperatures, hot and dry winds, and restricted soil water availability. These factors initiate a series of morphological, physiological, and biochemical disruptions that hinder wheat growth and productivity. Drought impact on wheat starts at biochemical levels through reactive oxygen species (ROS) generation and degradation of chlorophylls, and tolerance to stress is influenced by a polygenic system where numerous genes contribute minor effects and interact significantly with environmental factors transitioning to osmoprotectants. At the physiological level, drought alters the water content in the plant body, leading to reduced net photosynthetic rates, stomatal conductance, transpiration rates, and water utilization efficiency. At the morphological level, drought impacts all kinds of structures such as roots, shoots, leaves and reproductive parts. To counter these effects, wheat develops a set of tolerant mechanisms called drought escape, avoidance and tolerance. An increase in trichomes and leaf waxes, alteration of root\u2013shoot ratios, the staying green phenomenon, production of stress proteins like proline, activity of enzymes including superoxide dismutase (SOD), ascorbate peroxidase, catalase, etc., osmotic adjustment, abscisic acid (ABA) accumulation, expression of dehydration proteins called dehydrin, etc., contribute towards drought tolerance. This comprehensive review investigates the intricate interactions between drought and various wheat genotypes, emphasizing their substantial impacts on plant physiology, biochemistry, growth dynamics, and grain yield. Additionally, this review assesses a variety of genetic and biotechnological strategies aimed at enhancing the resilience of wheat genotypes to drought stress. By integrating recent research findings with practical applications, this review provides a detailed framework for improving the adaptive capacity of wheat plants to withstand the escalating threats of drought stress, thereby supporting sustainable wheat production in a changing climate. Addressing drought stress through genetic and biotechnological management practices is crucial for maintaining wheat productivity.<\/jats:p>","DOI":"10.3390\/plants15172653","type":"journal-article","created":{"date-parts":[[2026,8,31]],"date-time":"2026-08-31T02:08:58Z","timestamp":1788142138000},"page":"2653","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Wheat Drought Management: A Broader Prospect"],"prefix":"10.3390","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0009-0002-3164-7336","authenticated-orcid":false,"given":"Asfa","family":"Batool","sequence":"first","affiliation":[{"name":"Key Laboratories of Economic Forest Germplasm Improvement and Comprehensive Resources Utilization of Hubei Province, Collaborative Innovation Center for the Characteristic Resources Exploitation of Dabie Mountains, College of Biology and Agricultural Resources, Huanggang Normal University, Huanggang 438000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shi-Sheng","family":"Li","sequence":"additional","affiliation":[{"name":"Key Laboratories of Economic Forest Germplasm Improvement and Comprehensive Resources Utilization of Hubei Province, Collaborative Innovation Center for the Characteristic Resources Exploitation of Dabie Mountains, College of Biology and Agricultural Resources, Huanggang Normal University, Huanggang 438000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wei","family":"Tu","sequence":"additional","affiliation":[{"name":"Key Laboratories of Economic Forest Germplasm Improvement and Comprehensive Resources Utilization of Hubei Province, Collaborative Innovation Center for the Characteristic Resources Exploitation of Dabie Mountains, College of Biology and Agricultural Resources, Huanggang Normal University, Huanggang 438000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yun-Li","family":"Xiao","sequence":"additional","affiliation":[{"name":"Key Laboratories of Economic Forest Germplasm Improvement and Comprehensive Resources Utilization of Hubei Province, Collaborative Innovation Center for the Characteristic Resources Exploitation of Dabie Mountains, College of Biology and Agricultural Resources, Huanggang Normal University, Huanggang 438000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ting","family":"Zhou","sequence":"additional","affiliation":[{"name":"School of Agriculture and Biomanufacturing, Zhengzhou University, Zhengzhou 450001, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hongyuan","family":"Du","sequence":"additional","affiliation":[{"name":"Key Laboratories of Economic Forest Germplasm Improvement and Comprehensive Resources Utilization of Hubei Province, Collaborative Innovation Center for the Characteristic Resources Exploitation of Dabie Mountains, College of Biology and Agricultural Resources, Huanggang Normal University, Huanggang 438000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,8,29]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"1753","DOI":"10.1007\/s00122-021-03795-1","article-title":"Breeding for Drought and Heat Tolerance in Wheat","volume":"134","author":"Langridge","year":"2021","journal-title":"Theor. Appl. Genet."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Bapela, T., Shimelis, H., Tsilo, T., and Mathew, I. (2022). Genetic Improvement of Wheat for Drought Tolerance: Progress, Challenges and Opportunities. Plants, 11.","DOI":"10.3390\/plants11101331"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"212","DOI":"10.1007\/s10142-024-01488-8","article-title":"Drought-Tolerant Wheat for Enhancing Global Food Security","volume":"24","author":"Bohra","year":"2024","journal-title":"Funct. Integr. Genom."},{"key":"ref_4","first-page":"311","article-title":"Growth, Physiological Performance and Yield Traits Responses in Bread Wheat Cultivars Under Drought, Sprinkler Irrigation and Potassium Levels Conditions","volume":"15","author":"Ghanem","year":"2024","journal-title":"J. Plant Prod."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Singh, C., Yadav, S., Khare, V., Gupta, V., Patial, M., Kumar, S., Mishra, C., Tyagi, B., Gupta, A., and Sharma, A.K. (2025). Wheat Drought Tolerance: Unveiling a Synergistic Future with Conventional and Molecular Breeding Strategies. Plants, 14.","DOI":"10.3390\/plants14071053"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"2296094","DOI":"10.1080\/23311932.2023.2296094","article-title":"Drought Stress Effect, Tolerance, and Management in Wheat\u2014A Review","volume":"10","author":"Nyaupane","year":"2024","journal-title":"Cogent Food Agric."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Sallam, A., Alqudah, A., Dawood, M., Baenziger, P., and B\u00f6rner, A. (2019). Drought Stress Tolerance in Wheat and Barley: Advances in Physiology, Breeding and Genetics Research. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20133137"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"331","DOI":"10.1080\/07352689.2014.875291","article-title":"Drought Stress in Wheat during Flowering and Grain-Filling Periods","volume":"33","author":"Farooq","year":"2014","journal-title":"Crit. Rev. Plant Sci."},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Ahmad, A., Aslam, Z., Hussain, S., Javed, T., Hussain, S., Bashir, S., Hussain, I., Belliturk, K., Adamski, R., and Siuta, D. (2022). Soil Application of Wheat Straw Vermicompost Enhances Morpho-Physiological Attributes and Antioxidant Defense in Wheat Under Drought Stress. Front. Environ. Sci., 10.","DOI":"10.3389\/fenvs.2022.894517"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"17189","DOI":"10.1038\/s41598-020-74303-6","article-title":"Metabolic and Physiological Responses to Progressive Drought Stress in Bread Wheat","volume":"10","author":"Itam","year":"2020","journal-title":"Sci. Rep."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1080\/02571862.2025.2497229","article-title":"Managing Drought Stress in Wheat (Triticum aestivum L.) Production: Strategies and Impacts","volume":"42","author":"Mutanda","year":"2025","journal-title":"S. Afr. J. Plant Soil"},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Vassileva, V., Georgieva, M., Zehirov, G., and Dimitrova, A. (2023). Exploring the Genotype-Dependent Toolbox of Wheat Under Drought Stress. Agriculture, 13.","DOI":"10.20944\/preprints202308.0476.v1"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"80","DOI":"10.1007\/s11738-018-2651-6","article-title":"Physiological Responses of Wheat to Drought Stress and Its Mitigation Approaches","volume":"40","author":"Ahmad","year":"2018","journal-title":"Acta Physiol. Plant."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"164","DOI":"10.26832\/24566632.2025.1001024","article-title":"A Comprehensive Review on Drought Stress in Wheat: Causes, Mechanism and Management Practices","volume":"10","author":"Sharma","year":"2025","journal-title":"Arch. Agric. Environ. Sci."},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Wan, C., Dang, P., Gao, L., Wang, J., Tao, J., Qin, X., Feng, B., and Gao, J. (2022). How Does the Environment Affect Wheat Yield and Protein Content Response to Drought? A Meta-Analysis. Front. Plant Sci., 13.","DOI":"10.3389\/fpls.2022.896985"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"108683","DOI":"10.1016\/j.plaphy.2024.108683","article-title":"OsJAZ10 Negatively Modulates the Drought Tolerance by Integrating Hormone Signaling with Systemic Electrical Activity in Rice","volume":"211","author":"Wu","year":"2024","journal-title":"Plant Physiol. Biochem."},{"key":"ref_17","first-page":"3","article-title":"Drought Stress and Its Management in Wheat (Triticum aestivum L.): A Review","volume":"14","author":"Sharma","year":"2022","journal-title":"Agric. Sci. Technol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"104517","DOI":"10.1016\/j.envexpbot.2021.104517","article-title":"Grain Development in Wheat Under Combined Heat and Drought Stress: Plant Responses and Management","volume":"188","author":"Zahra","year":"2021","journal-title":"Environ. Exp. Bot."},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"Khan, A., Wang, Y.-F., Akbar, R., and Alhoqail, W. (2025). Mechanistic Insights and Future Perspectives of Drought Stress Management in Staple Crops. Front. Plant Sci., 16.","DOI":"10.3389\/fpls.2025.1547452"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"26","DOI":"10.3390\/stresses2010003","article-title":"Response of Wheat Genotypes to Drought Stress Stimulated by PEG","volume":"2","author":"Sharma","year":"2022","journal-title":"Stresses"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"78","DOI":"10.54386\/jam.v16i1.1490","article-title":"Identification of Physiological and Yield Related Traits of Wheat (Triticum aestivum L.) Under Varying Soil Moisture Stress","volume":"16","author":"Sharma","year":"2022","journal-title":"J. Agrometeorol."},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Wu, H., and Yang, Z. (2024). Effects of Drought Stress and Postdrought Rewatering on Winter Wheat: A Meta-Analysis. Agronomy, 14.","DOI":"10.3390\/agronomy14020298"},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Khan, N. (2025). Molecular Insights into ABA-Mediated Regulation of Stress Tolerance and Development in Plants. Int. J. Mol. Sci., 26.","DOI":"10.3390\/ijms26167872"},{"key":"ref_24","first-page":"702","article-title":"Molecular and Eco-Physiological Responses of Wheat (Triticum aestivum L.) to Drought Stress: Implications for Climate-Resilient Crop Improvement","volume":"4","author":"Babar","year":"2026","journal-title":"Phys. Educ. Health Soc. Sci."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"16","DOI":"10.3844\/ajabssp.2024.16.22","article-title":"Drought and Terminal Heat Stress Effects and Its Management on Wheat Genotypes: A Review","volume":"19","author":"Gupta","year":"2024","journal-title":"Am. J. Agric. Biol. Sci."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"67","DOI":"10.26480\/mjsa.02.2021.67.76","article-title":"Drought Stress Impacts on Wheat and Its Resistance Mechanisms","volume":"5","author":"Rijal","year":"2020","journal-title":"Malays. J. Sustain. Agric."},{"key":"ref_27","doi-asserted-by":"crossref","unstructured":"Nagy, D., Meszl\u00e9nyi, T., Boda, K., Lantos, C., and Pauk, J. (2025). Agronomical Responses of Elite Winter Wheat (Triticum aestivum L.) Varieties in Phenotyping Experiments Under Continuous Water Withdrawal and Optimal Water Management in Greenhouses. Plants, 14.","DOI":"10.3390\/plants14152435"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"529","DOI":"10.9734\/ijecc\/2024\/v14i13866","article-title":"Drought Stress and Its Tolerance Mechanism in Wheat","volume":"14","author":"Gupta","year":"2024","journal-title":"Int. J. Environ. Clim. Chang."},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Khamis, G., Alsherif, E., Korany, S., Aldailami, D., Aloufi, A., Khalaf, M., Hamed, S., Maridue\u00f1a-Zavala, M., Jaouni, S.A., and Selim, S. (2025). Drought Stress Differentially Influences Growth, Physiology, and Metabolite Accumulation in Triticum aestivum L. (C3) and Amaranthus caudatus L. (C4) Plants. BMC Plant Biol., 25.","DOI":"10.1186\/s12870-025-07022-7"},{"key":"ref_30","doi-asserted-by":"crossref","unstructured":"Wasaya, A., Manzoor, S., Yasir, T., Sarwar, N., Mubeen, K., Ismail, I., Raza, A., Hossain, A., and Sabagh, E. (2021). Evaluation of Fourteen Bread Wheat (Triticum aestivum L.) Genotypes by Observing Gas Exchange Parameters, Relative Water and Chlorophyll Content, and Yield Attributes Under Drought Stress. Sustainability, 13.","DOI":"10.3390\/su13094799"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"2836","DOI":"10.1111\/1365-2745.13988","article-title":"Elevated CO2 Alleviates Adverse Effects of Drought on Plant Water Relations and Photosynthesis: A Global Meta-analysis","volume":"110","author":"Wang","year":"2022","journal-title":"J. Ecol."},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Guizani, A., Askri, H., Amenta, M.L., Defez, R., Babay, E., Bianco, C., Rapan\u00e1, N., Finetti-Sialer, M., and Gharbi, F. (2023). Drought Responsiveness in Six Wheat Genotypes: Identification of Stress Resistance Indicators. Front. Plant Sci., 14.","DOI":"10.3389\/fpls.2023.1232583"},{"key":"ref_33","doi-asserted-by":"crossref","unstructured":"Chowdhury, M., Hasan, M., Bahadur, M., Islam, M., Hakim, M., Iqbal, M., Javed, T., Raza, A., Shabbir, R., and Sorour, S. (2021). Evaluation of Drought Tolerance of Some Wheat (Triticum aestivum L.) Genotypes Through Phenology, Growth, and Physiological Indices. Agronomy, 11.","DOI":"10.3390\/agronomy11091792"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"5238","DOI":"10.1016\/j.sjbs.2021.05.040","article-title":"Improvement in Drought Tolerance in Bread Wheat Is Related to an Improvement in Osmolyte Production, Antioxidant Enzyme Activities, and Gaseous Exchange","volume":"28","author":"Qayyum","year":"2021","journal-title":"Saudi J. Biol. Sci."},{"key":"ref_35","doi-asserted-by":"crossref","unstructured":"Pecka, J., Kraus, K., Zelen\u00fd, M., and Hnili\u010dkov\u00e1, H. (2025). Exogenous Proline Modulates Physiological Responses and Induces Stress Memory in Wheat Under Repeated and Delayed Drought Stress. Agronomy, 15.","DOI":"10.3390\/agronomy15061370"},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Qiao, M., Hong, C., Jiao, Y., Hou, S., and Gao, H. (2024). Impacts of Drought on Photosynthesis in Major Food Crops and the Related Mechanisms of Plant Responses to Drought. Plants, 13.","DOI":"10.3390\/plants13131808"},{"key":"ref_37","first-page":"279","article-title":"Silicon Improves Root System and Canopy Physiology in Wheat Under Drought Stress","volume":"502","author":"Ashfaq","year":"2023","journal-title":"Plant Soil"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"645","DOI":"10.1007\/s11033-025-10742-6","article-title":"Physiological and Molecular Responses of Bread Wheat and Its Wild Relative Species to Drought Stress","volume":"52","author":"Jadidi","year":"2025","journal-title":"Mol. Biol. Rep."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Popova, A., Mihailova, G., Geneva, M., Peeva, V., Kirova, E., Sichanova, M., Dobrikova, A., and Georgieva, K. (2023). Different Responses to Water Deficit of Two Common Winter Wheat Varieties: Physiological and Biochemical Characteristics. Plants, 12.","DOI":"10.3390\/plants12122239"},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Wasaya, A., Affan, M., Yasir, T.A., Mubeen, K., Rehman, H., Ali, M., Nawaz, F., Galal, A., and Iqbal, M. (2021). Foliar Potassium Sulfate Application Improved Photosynthetic Characteristics, Water Relations and Seedling Growth of Drought-Stressed Maize. Atmosphere, 12.","DOI":"10.3390\/atmos12060663"},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Qiao, L., Chang, L., Kai, M., Zhang, X., Kang, T., Wu, L., Zhang, X., Li, X., Zhao, J., and Zhao, Z. (2024). Exploring Drought Resistance Genes from the Roots of the Wheat Cultivar Yunhan1818. Int. J. Mol. Sci., 25.","DOI":"10.3390\/ijms252413458"},{"key":"ref_42","doi-asserted-by":"crossref","unstructured":"Wang, J., Zhang, X., Han, Z., Feng, H., Wang, Y., Kang, J., Han, X., Wang, L., Wang, C., and Li, H. (2022). Analysis of Physiological Indicators Associated with Drought Tolerance in Wheat Under Drought and Re-Watering Conditions. Antioxidants, 11.","DOI":"10.3390\/antiox11112266"},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Ahmad, A., Aslam, Z., Javed, T., Hussain, S., Raza, A., Shabbir, R., Mora-Poblete, F., Saeed, T., Zulfiqar, F., and Ali, M. (2022). Screening of Wheat (Triticum aestivum L.) Genotypes for Drought Tolerance Through Agronomic and Physiological Response. Agronomy, 12.","DOI":"10.3390\/agronomy12020287"},{"key":"ref_44","doi-asserted-by":"crossref","unstructured":"Miranda, V., Silva-Castro, G., Ruiz-Lozano, J., Fracchia, S., and Garc\u00eda-Romera, I. (2023). Fungal Endophytes Enhance Wheat and Tomato Drought Tolerance in Terms of Plant Growth and Biochemical Parameters. J. Fungi, 9.","DOI":"10.3390\/jof9030384"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"e05908","DOI":"10.1016\/j.heliyon.2021.e05908","article-title":"Salicylic Acid Alleviated the Effect of Drought Stress on Photosynthetic Characteristics and Leaf Protein Pattern in Winter Wheat","volume":"7","author":"Khalvandi","year":"2021","journal-title":"Heliyon"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"e70103","DOI":"10.1002\/fes3.70103","article-title":"Melatonin and L-Cysteine Desulfhydrase: Unraveling Hydrogen Sulfide Signaling for Drought Tolerance in Bread Wheat (Triticum aestivum)","volume":"14","author":"Muslemyar","year":"2025","journal-title":"Food Energy Secur."},{"key":"ref_47","doi-asserted-by":"crossref","unstructured":"Todorova, D., Sergiev, I., Katerova, Z., Shopova, E., Dimitrova, L., and Brankova, L. (2021). Assessment of the Biochemical Responses of Wheat Seedlings to Soil Drought after Application of Selective Herbicide. Plants, 10.","DOI":"10.3390\/plants10040733"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"696","DOI":"10.1111\/ppl.13237","article-title":"Effect of 24-Epibrassinolide on Plant Growth, Antioxidants Defense System and Endogenous Hormones in Two Wheat Varieties Under Drought Stress","volume":"172","author":"Khan","year":"2020","journal-title":"Physiol. Plant."},{"key":"ref_49","doi-asserted-by":"crossref","unstructured":"Moloi, S., Alqarni, A., Brown, A., Goche, T., Shargie, N., Moloi, M., Gokul, A., Chivasa, S., and Ngara, R. (2024). Comparative Physiological, Biochemical, and Leaf Proteome Responses of Contrasting Wheat Varieties to Drought Stress. Plants, 13.","DOI":"10.3390\/plants13192797"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"440","DOI":"10.1007\/s13205-021-02991-6","article-title":"Effect of Progressive Drought Stress on Physio-Biochemical Responses and Gene Expression Patterns in Wheat","volume":"11","author":"Amoah","year":"2021","journal-title":"3 Biotech"},{"key":"ref_51","doi-asserted-by":"crossref","unstructured":"Duvnjak, J., \u0160ar\u010devi\u0107, H., Vukovi\u0107, R., and \u0160pani\u0107, V. (2024). Effects of Drought at Anthesis on Flag Leaf Physiology and Gene Expression in Diverse Wheat (Triticum aestivum L.) Genotypes. Agronomy, 14.","DOI":"10.3390\/agronomy14071522"},{"key":"ref_52","doi-asserted-by":"crossref","unstructured":"Shan, H., Wen, H., Zhang, J., Wang, Y., Lu, L., Liu, Y., Yang, B., and Ji, W. (2025). Synergistic Role of Streptomyces Composite Inoculants in Mitigating Wheat Drought Stress Under Field Conditions. Plants, 14.","DOI":"10.3390\/plants14030366"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"15985","DOI":"10.1038\/s41598-024-66506-y","article-title":"Mitigating Drought-Induced Oxidative Stress in Wheat (Triticum aestivum L.) Through Foliar Application of Sulfhydryl Thiourea","volume":"14","author":"Ishfaq","year":"2024","journal-title":"Sci. Rep."},{"key":"ref_54","doi-asserted-by":"crossref","unstructured":"Isgandarova, T., Rustamova, S., Aliyeva, D., Rzayev, F., Gasimov, E., and Huseynova, I. (2024). Antioxidant and Ultrastructural Alterations in Wheat During Drought-Induced Leaf Senescence. Agronomy, 14.","DOI":"10.20944\/preprints202410.0728.v1"},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Lamlom, S., Abdelghany, A., Farouk, A., Alwakel, E., Makled, K., Bukhari, N., Hatamleh, A., Ren, H., El-Sorady, G., and Shehab, A. (2025). Biochemical and Yield Response of Spring Wheat to Drought Stress Through Gibberellic and Abscisic Acids. BMC Plant Biol., 25.","DOI":"10.1186\/s12870-024-05879-8"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"591","DOI":"10.1007\/s42976-023-00414-1","article-title":"Wheat (Triticum aestivum L.) Biochemical and Nutritional Properties Affected by Plant Growth Regulators Under Field Drought Conditions","volume":"52","author":"Azizi","year":"2023","journal-title":"Cereal Res. Commun."},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Hussein, H., Alshammari, S., El-Sadek, M.A., Kenawy, S., and Badawy, A. (2023). The Promotive Effect of Putrescine on Growth, Biochemical Constituents, and Yield of Wheat (Triticum aestivum L.) Plants Under Water Stress. Agriculture, 13.","DOI":"10.3390\/agriculture13030587"},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Licaj, I., Felice, D., Germinario, C., Zanotti, C., Fiorillo, A., Marra, M., and Rocco, M. (2023). An Artificial Intelligence-Integrated Analysis of the Effect of Drought Stress on Root Traits of \u201cModern\u201d and \u201cAncient\u201d Wheat Varieties. Front. Plant Sci., 14.","DOI":"10.3389\/fpls.2023.1241281"},{"key":"ref_59","first-page":"112906","article-title":"Dynamic Structural Changes in Wheat Seedlings as a Response to Drought Stress","volume":"365","author":"Leszczuk","year":"2025","journal-title":"Plant Sci. Int. J. Exp. Plant Biol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"61","DOI":"10.36320\/ajb\/v14.i3.11164","article-title":"Some Biochemical and Biomass Responses of Wheat [Triticum aestivum L.] to Suboptimal Water Supply and Different Potassium Rates","volume":"14","author":"Hussein","year":"2023","journal-title":"AL-Kufa Univ. J. Biol."},{"key":"ref_61","doi-asserted-by":"crossref","unstructured":"Ullah, S., Khan, M., Lodhi, S., Ahmed, I., Tayyab, M., Mehmood, T., Din, I., Khan, M., Sohail, Q., and Akram, M. (2022). Targeted Metabolomics Reveals Fatty Acid Abundance Adjustments as Playing a Crucial Role in Drought-Stress Response and Post-Drought Recovery in Wheat. Front. Genet., 13.","DOI":"10.3389\/fgene.2022.972696"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"100708","DOI":"10.1016\/j.wace.2024.100708","article-title":"Probabilistic Analysis of Drought Impact on Wheat Yield and Climate Change Implications","volume":"45","author":"Wang","year":"2024","journal-title":"Weather Clim. Extrem."},{"key":"ref_63","doi-asserted-by":"crossref","unstructured":"Seleiman, M., Al-Suhaibani, N., Ali, N., Akmal, M., Alotaibi, M., Refay, Y., Dindaro\u011flu, T., Abdul-Wajid, H.H., and Battaglia, M. (2021). Drought Stress Impacts on Plants and Different Approaches to Alleviate Its Adverse Effects. Plants, 10.","DOI":"10.3390\/plants10020259"},{"key":"ref_64","doi-asserted-by":"crossref","unstructured":"Pour-Aboughadareh, A., Mohammadi, R., Etminan, A., Shooshtari, L., Maleki-Tabrizi, N., and Poczai, P. (2020). Effects of Drought Stress on Some Agronomic and Morpho-Physiological Traits in Durum Wheat Genotypes. Sustainability, 12.","DOI":"10.3390\/su12145610"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"1291","DOI":"10.1111\/ppl.13423","article-title":"Drought: Sensing, Signalling, Effects and Tolerance in Higher Plants","volume":"172","author":"Mukarram","year":"2021","journal-title":"Physiol. Plant."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"51","DOI":"10.47115\/bsagriculture.1573391","article-title":"Effect of Potassium Optimization on Wheat Drought Tolerance in Controlled Conditions","volume":"8","year":"2025","journal-title":"Black Sea J. Agric."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"1239","DOI":"10.1093\/aob\/mcaf152","article-title":"Root Morphology and Anatomy Respond Similarly to Drought and Flooding in Two Wheat Cultivars","volume":"136","author":"Guhr","year":"2025","journal-title":"Ann. Bot."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"e12015","DOI":"10.7717\/peerj.12015","article-title":"The Relationship Between Characteristics of Root Morphology and Grain Filling in Wheat Under Drought Stress","volume":"9","author":"Chen","year":"2021","journal-title":"PeerJ"},{"key":"ref_69","doi-asserted-by":"crossref","unstructured":"Wang, Q., Wu, Y., Ozavize, S.F., Qiu, C., Holford, P., and Wu, F.-B. (2024). Genotypic Differences in Morphological, Physiological and Agronomic Traits in Wheat (Triticum aestivum L.) in Response to Drought. Plants, 13.","DOI":"10.3390\/plants13020307"},{"key":"ref_70","doi-asserted-by":"crossref","unstructured":"Chen, M., Zhu, X., Liu, X., Wu, C., Yu, C., Hu, G., Chen, L., Chen, R., Bouzayen, M., and Zouine, M. (2021). Knockout of Auxin Response Factor SlARF4 Improves Tomato Resistance to Water Deficit. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22073347"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"110697","DOI":"10.1016\/j.plaphy.2025.110697","article-title":"The LBD Transcription Factor TaLBD16 Positively Regulates Drought Tolerance in Wheat","volume":"229","author":"Wang","year":"2025","journal-title":"Plant Physiol. Biochem."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"1344","DOI":"10.1093\/plphys\/kiac523","article-title":"Wheat TaSnRK2.10 Phosphorylates TaERD15 and TaENO1 and Confers Drought Tolerance When Overexpressed in Rice","volume":"191","author":"Zhang","year":"2022","journal-title":"Plant Physiol."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"527","DOI":"10.1007\/s11033-024-09481-x","article-title":"An SNP Based Genotyping Assay for Genes Associated with Drought Tolerance in Bread Wheat","volume":"51","author":"Rauf","year":"2024","journal-title":"Mol. Biol. Rep."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"352","DOI":"10.1111\/pbi.14501","article-title":"Wheat TaPYL9-involved Signalling Pathway Impacts Plant Drought Response Through Regulating Distinct Osmotic Stress-Associated Physiological Indices","volume":"23","author":"Zhang","year":"2024","journal-title":"Plant Biotechnol. J."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1016\/j.plaphy.2026.111397","article-title":"Functional Characterization of TaSnRK2.8-5A Reveals the Central Role of Its Signal Module in Enhancing Drought Tolerance Through Coordinated Molecular and Physiological Response of Wheat","volume":"235","author":"Liu","year":"2026","journal-title":"Plant Physiol. Biochem."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"683","DOI":"10.1093\/jxb\/erp331","article-title":"TaSnRK2.4, an SNF1-Type Serine\/Threonine Protein Kinase of Wheat (Triticum aestivum L.), Confers Enhanced Multistress Tolerance in Arabidopsis","volume":"61","author":"Mao","year":"2009","journal-title":"J. Exp. Bot."},{"key":"ref_77","doi-asserted-by":"crossref","unstructured":"Zhang, M., Du, G., Zhang, H., Zheng, R., Zhao, L., Huang, M., Wang, X., Liu, K., and Yan, D. (2026). Metabolomics and Biochemical Analysis Reveal the Regulatory Mechanism of Exogenous Sorbitol-Chelated Potassium on Wheat Under Drought Stress. Front. Plant Sci., 17.","DOI":"10.3389\/fpls.2026.1751075"},{"key":"ref_78","first-page":"6440","article-title":"TaSnRK2.1-2D Contributes to Drought Tolerance by Modulating ROS Production in Wheat","volume":"48","author":"Bhanbhro","year":"2025","journal-title":"Plant"},{"key":"ref_79","first-page":"79","article-title":"Functional Characterization on Wheat Protein Kinase Gene TaSnRK2.1 in Mediating Plant Resistance to Drought Stress","volume":"2","author":"Chenyang","year":"2023","journal-title":"Life Sci. Technol."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"345","DOI":"10.1016\/j.cj.2025.10.017","article-title":"Phosphatase Member TaPP2C62-2A Cooperates with Downstream Partners to Regulate Drought Response in Wheat","volume":"14","author":"Hou","year":"2025","journal-title":"Crop J."},{"key":"ref_81","doi-asserted-by":"crossref","unstructured":"Zhang, H., Mao, X., Wang, C., and Jing, R. (2010). Overexpression of a Common Wheat Gene TaSnRK2.8 Enhances Tolerance to Drought, Salt and Low Temperature in Arabidopsis. PLoS ONE, 5.","DOI":"10.1371\/journal.pone.0016041"},{"key":"ref_82","doi-asserted-by":"crossref","unstructured":"Dong, F., Liu, Y., Zhang, H., Li, Y., Song, J., Chen, S., Wang, S., Zhu, Z., Li, Y., and Liu, Y. (2025). TaSnRK3.23B, a CBL-Interacting Protein Kinase of Wheat, Confers Drought Stress Tolerance by Promoting ROS Scavenging in Arabidopsis. BMC Plant Biol., 25.","DOI":"10.1186\/s12870-025-06091-y"},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"83","DOI":"10.18524\/2077-1746.2020.2(47).218456","article-title":"Approbation of TaSnRK2.8-A Gene Marker Analysis on Bread Winter Wheat Varieties","volume":"25","author":"Chebotar","year":"2020","journal-title":"Odesa Natl. Univ. Her. Biol."},{"key":"ref_84","doi-asserted-by":"crossref","unstructured":"Cevher-Keskin, B., Y\u0131ld\u0131zhan, Y., Sekmen, A., Fayetorbay, R., Sezerman, O., \u00d6zer, B., Onar\u0131c\u0131, S., T\u00fcrkan, \u0130., and T\u00f6r, M. (2025). Deciphering Drought-Response in Wheat (Triticum aestivum): Physiological, Biochemical, and Transcriptomic Insights into Tolerant and Sensitive Cultivars Under Dehydration Shock. Front. Plant Sci., 16.","DOI":"10.3389\/fpls.2025.1649378"},{"key":"ref_85","doi-asserted-by":"crossref","unstructured":"Kulkarni, M., Soolanayakanahally, R., Ogawa, S., Uga, Y., Selvaraj, M., and Kagale, S. (2017). Drought Response in Wheat: Key Genes and Regulatory Mechanisms Controlling Root System Architecture and Transpiration Efficiency. Front. Chem., 5.","DOI":"10.3389\/fchem.2017.00106"},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"1565","DOI":"10.1111\/pbi.14605","article-title":"A System Genetics Analysis Uncovers the Regulatory Variants Controlling Drought Response in Wheat","volume":"23","author":"Chen","year":"2025","journal-title":"Plant Biotechnol. J."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"109086","DOI":"10.1016\/j.plaphy.2024.109086","article-title":"The Transcription Factor TabZIP156 Acts as a Positive Regulator in Response to Drought Tolerance in Arabidopsis and Wheat (Triticum aestivum L.)","volume":"216","author":"Bu","year":"2024","journal-title":"Plant Physiol. Biochem."},{"key":"ref_88","doi-asserted-by":"crossref","unstructured":"Nouraei, S., Mia, M.S., Liu, H., Turner, N., and Yan, G. (2022). Transcriptome Analyses of Near Isogenic Lines Reveal Putative Drought Tolerance Controlling Genes in Wheat. Front. Plant Sci., 13.","DOI":"10.3389\/fpls.2022.857829"},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"6563","DOI":"10.1038\/s41467-025-61943-3","article-title":"An Elite Allele TaDT1-AhapI Enhances Drought Tolerance via Mediating Autophagic Pathways in Wheat","volume":"16","author":"Liu","year":"2025","journal-title":"Nat. Commun."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"100298","DOI":"10.1016\/j.stress.2023.100298","article-title":"Revisiting the Molecular Mechanisms and Adaptive Strategies Associated with Drought Stress Tolerance in Common Wheat (Triticum aestivum L.)","volume":"11","author":"Bhanbhro","year":"2023","journal-title":"Plant Stress"},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"210","DOI":"10.1111\/j.1399-3054.2011.01539.x","article-title":"Molecular Characterization of Novel TaNAC Genes in Wheat and Overexpression of TaNAC2a Confers Drought Tolerance in Tobacco","volume":"144","author":"Tang","year":"2012","journal-title":"Physiol. Plant."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"110578","DOI":"10.1016\/j.plaphy.2025.110578","article-title":"Functional Characterization of TaNAC6-3B: A Key Regulator of Drought Tolerance in Wheat (Triticum aestivum L.)","volume":"229","author":"Chen","year":"2025","journal-title":"Plant Physiol. Biochem."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"846","DOI":"10.1111\/pbi.13764","article-title":"The Wheat ABA Receptor Gene TaPYL1-1B Contributes to Drought Tolerance and Grain Yield by Increasing Water-use Efficiency","volume":"20","author":"Mao","year":"2021","journal-title":"Plant Biotechnol. J."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"847","DOI":"10.1111\/ppl.13268","article-title":"Transcription Factors as Key Molecular Target to Strengthen the Drought Stress Tolerance in Plants","volume":"172","author":"Manna","year":"2020","journal-title":"Physiol. Plant."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"2019","DOI":"10.1007\/s00122-016-2794-z","article-title":"Transcription Factors Involved in Drought Tolerance and Their Possible Role in Developing Drought Tolerant Cultivars with Emphasis on Wheat (Triticum aestivum L.)","volume":"129","author":"Gahlaut","year":"2016","journal-title":"Theor. Appl. Genet."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"230","DOI":"10.1111\/j.1467-7652.2010.00547.x","article-title":"Improvement of Stress Tolerance of Wheat and Barley by Modulation of Expression of DREB\/CBF Factors","volume":"9","author":"Morran","year":"2011","journal-title":"Plant Biotechnol. J."},{"key":"ref_97","doi-asserted-by":"crossref","unstructured":"Ge, M., Tang, Y., Guan, Y., Lv, M., Zhou, C., Ma, H., and Lv, J. (2024). TaWRKY31, a Novel WRKY Transcription Factor in Wheat, Participates in Regulation of Plant Drought Stress Tolerance. BMC Plant Biol., 24.","DOI":"10.1186\/s12870-023-04709-7"},{"key":"ref_98","doi-asserted-by":"crossref","unstructured":"Zhang, Y., Tu, M., Han, J., Zhang, Q., Li, L., Shi, F., Zhao, H., Yu, P., Du, C., and Wang, B. (J. Adv. Res., 2026). A Poaceae-Specific DREB-A6 Transcription Factor TaDREB26-B Regulates ABA Responses and Enhances Drought Stress Tolerance in Wheat, J. Adv. Res., in press.","DOI":"10.1016\/j.jare.2026.03.057"},{"key":"ref_99","doi-asserted-by":"crossref","unstructured":"Komal, L., Kamran, A., Jahan, S., Akram, W., Siddiqui, M., Alamri, S., and Shakir, M. (2025). Molecular Insights into Drought Tolerance in Wheat Through In-Silico Genome-Wide Analysis of DREB1 Transcription Factor and Peroxidase Interactions. BMC Plant Biol., 25.","DOI":"10.1186\/s12870-025-06938-4"},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"100058","DOI":"10.1016\/j.abiote.2026.100058","article-title":"A TaMYB2\u2013TaMAP3K17 Module Enhances Drought Tolerance by Promoting Reactive Oxygen Species Scavenging in Wheat","volume":"7","author":"Yang","year":"2026","journal-title":"aBIOTECH"},{"key":"ref_101","doi-asserted-by":"crossref","unstructured":"Zhao, X., Bai, S., Li, L., Han, X., Li, J., Zhu, Y., Fang, Y., Zhang, D., and Li, S. (2020). Comparative Transcriptome Analysis of Two Aegilops tauschii with Contrasting Drought Tolerance by RNA-Seq. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21103595"},{"key":"ref_102","doi-asserted-by":"crossref","unstructured":"Wang, Y., Jin, H., Du, S., Fang, B.-T., Yue, J.-Q., Yang, C., Wang, H.-F., Zhang, D., Wang, J., and Song, H. (2025). Multi-Omics Association Analysis of DOF Transcription Factors Involved in the Drought Resistance of Wheat Induced by Strigolactone. Int. J. Mol. Sci., 26.","DOI":"10.3390\/ijms26062396"},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"101379","DOI":"10.1016\/j.stress.2026.101379","article-title":"TaPHL7\u2013TaSOD7-4A Regulatory Pair Enhances Drought Tolerance in Wheat (Triticum aestivum L.)","volume":"21","author":"Yu","year":"2026","journal-title":"Plant Stress"},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"143008","DOI":"10.1016\/j.ijbiomac.2025.143008","article-title":"The Drought-Responsive Wheat AP2\/ERF Transcription Factor TaRAP2-13L and Its Interacting Protein TaWRKY10 Enhance Drought Tolerance in Transgenic Arabidopsis and Wheat (Triticum aestivum L.)","volume":"309","author":"Shao","year":"2025","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_105","first-page":"6524","article-title":"A Raf-Like MAPKKK Gene TaHT1 Controls Drought Tolerance and Primary Root Length in Wheat","volume":"48","author":"Wang","year":"2025","journal-title":"Plant"},{"key":"ref_106","doi-asserted-by":"crossref","unstructured":"Aslam, M.M., Waseem, M., Jakada, B., Okal, E., Lei, Z., Saqib, H., Yuan, W., Xu, W., and Zhang, Q. (2022). Mechanisms of Abscisic Acid-Mediated Drought Stress Responses in Plants. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms23031084"},{"key":"ref_107","doi-asserted-by":"crossref","unstructured":"Ali, S., Hayat, K., Iqbal, A., and Xie, L. (2020). Implications of Abscisic Acid in the Drought Stress Tolerance of Plants. Agronomy, 10.","DOI":"10.3390\/agronomy10091323"},{"key":"ref_108","doi-asserted-by":"crossref","unstructured":"Wang, J., Song, L., Gong, X., Xu, J., and Li, M.-H. (2020). Functions of Jasmonic Acid in Plant Regulation and Response to Abiotic Stress. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21041446"},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"120","DOI":"10.1016\/j.cj.2020.06.002","article-title":"Abscisic Acid and Jasmonic Acid Are Involved in Drought Priming-Induced Tolerance to Drought in Wheat","volume":"9","author":"Wang","year":"2020","journal-title":"Crop J."},{"key":"ref_110","doi-asserted-by":"crossref","unstructured":"Duvnjak, J., Lon\u010dari\u0107, A., Brklja\u010di\u0107, L., \u0160amec, D., \u0160ar\u010devi\u0107, H., Salopek-Sondi, B., and \u0160pani\u0107, V. (2023). Morpho-Physiological and Hormonal Response of Winter Wheat Varieties to Drought Stress at Stem Elongation and Anthesis Stages. Plants, 12.","DOI":"10.3390\/plants12030418"},{"key":"ref_111","doi-asserted-by":"crossref","unstructured":"Saradadevi, R., Palta, J., and Siddique, K. (2017). ABA-Mediated Stomatal Response in Regulating Water Use during the Development of Terminal Drought in Wheat. Front. Plant Sci., 8.","DOI":"10.3389\/fpls.2017.01251"},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"1545","DOI":"10.30848\/PJB2021-5(11)","article-title":"Interactive Effects of Abscisic Acid (ABA) and Drought Stress on the Physiological Responses of Winter Wheat (Triticum aestivum L.)","volume":"53","author":"Kong","year":"2021","journal-title":"Pak. J. Bot."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"20411","DOI":"10.1038\/s41598-024-71404-4","article-title":"Abscisic Acid Improves Drought Resilience, Growth, Physio-Biochemical and Quality Attributes in Wheat (Triticum aestivum L.) at Critical Growth Stages","volume":"14","author":"Zulfiqar","year":"2024","journal-title":"Sci. Rep."},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"1209","DOI":"10.1007\/s10725-024-01190-6","article-title":"Foliar Application of Abscisic Acid and Glycine Betaine Induces Tolerance to Water Scarcity in Wheat","volume":"104","author":"Haque","year":"2024","journal-title":"Plant Growth Regul."},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"e30598","DOI":"10.1016\/j.heliyon.2024.e30598","article-title":"Abscisic Acid and Glycine Betaine-Mediated Seed and Root Priming Enhance Seedling Growth and Antioxidative Defense in Wheat Under Drought","volume":"10","author":"Baroi","year":"2024","journal-title":"Heliyon"},{"key":"ref_116","doi-asserted-by":"crossref","unstructured":"Li, M., Yang, Y., Wu, Y., Zheng, X., Zhao, J., Wu, B., Xu, Y., Zhang, G., Zhao, Z., and Wang, J. (2026). Jasmonates Regulate Drought Resistance by Controlling the Secondary Metabolism in Wheat Seedlings. BMC Plant Biol., 26.","DOI":"10.1186\/s12870-026-08574-y"},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"809","DOI":"10.1111\/ppl.13247","article-title":"Exogenous Abscisic Acid and Jasmonic Acid Restrain PEG-Induced Drought by Improving the Growth and Antioxidative Enzyme Activities in Pearl Millet","volume":"172","author":"Awan","year":"2020","journal-title":"Physiol. Plant."},{"key":"ref_118","doi-asserted-by":"crossref","unstructured":"Rasheed, A., Li, H., Tahir, M., Mahmood, A., Nawaz, M., Shah, A., Aslam, M., Negm, S., Moustafa, M., and Hassan, M. (2022). The Role of Nanoparticles in Plant Biochemical, Physiological, and Molecular Responses Under Drought Stress: A Review. Front. Plant Sci., 13.","DOI":"10.3389\/fpls.2022.976179"},{"key":"ref_119","doi-asserted-by":"crossref","first-page":"109550","DOI":"10.1016\/j.agwat.2025.109550","article-title":"Harnessing Nanoparticles to Enhance Crop Production under Drought Stress: A Quantitative Meta-Analysis","volume":"315","author":"Li","year":"2025","journal-title":"Agric. Water Manag."},{"key":"ref_120","doi-asserted-by":"crossref","first-page":"2700","DOI":"10.1038\/s41598-023-29784-6","article-title":"Morpho-Physiological and Biochemical Response of Wheat to Various Treatments of Silicon Nano-Particles Under Drought Stress Conditions","volume":"13","author":"Raza","year":"2023","journal-title":"Sci. Rep."},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"905","DOI":"10.1071\/FP21079","article-title":"Seed Nanopriming by Silicon Oxide Improves Drought Stress Alleviation Potential in Wheat Plants","volume":"48","author":"Tomar","year":"2021","journal-title":"Funct. Plant Biol."},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"5309","DOI":"10.1038\/s41598-025-89718-2","article-title":"ZnO-Nanoparticles and Stage-Based Drought Tolerance in Wheat (Triticum aestivum L.): Effect on Morpho-Physiology, Nutrients Uptake, Grain Yield and Quality","volume":"15","author":"Raza","year":"2025","journal-title":"Sci. Rep."},{"key":"ref_123","doi-asserted-by":"crossref","first-page":"2343","DOI":"10.1007\/s42976-025-00688-7","article-title":"Nanopriming as an Approach to Induce Tolerance against Drought Stress in Wheat Cultivars","volume":"53","author":"Megahed","year":"2025","journal-title":"Cereal Res. Commun."},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"11612","DOI":"10.1038\/s41598-026-43113-7","article-title":"Quantitative Trait Loci Associated with Drought Stress Tolerance in Wheat Primed with Zinc Oxide Nanoparticles at Seed Germination and Seedling Stages","volume":"16","author":"Mahmoud","year":"2026","journal-title":"Sci. Rep."},{"key":"ref_125","doi-asserted-by":"crossref","unstructured":"Omar, A., Heikal, Y., Zayed, E., Shamseldin, S., Salama, Y., Amer, K., Basuoni, M., Ellatif, S.A., and Mohamed, A. (2023). Conferring of Drought and Heat Stress Tolerance in Wheat (Triticum aestivum L.) Genotypes and Their Response to Selenium Nanoparticles Application. Nanomaterials, 13.","DOI":"10.3390\/nano13060998"},{"key":"ref_126","doi-asserted-by":"crossref","unstructured":"Hasanuzzaman, M., Raihan, M., Siddika, A., Bardhan, K., Hosen, M.S., and Prasad, P. (2024). Selenium and Its Nanoparticles Modulate the Metabolism of Reactive Oxygen Species and Morpho-Physiology of Wheat (Triticum aestivum L.) to Combat Oxidative Stress Under Water Deficit Conditions. BMC Plant Biol., 24.","DOI":"10.1186\/s12870-024-05282-3"},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1186\/s40538-025-00778-y","article-title":"Silver Nanoparticles Priming for Drought Tolerance in Wheat: Insights from Antioxidant System Activation and Stress Memory","volume":"12","author":"Ding","year":"2025","journal-title":"Chem. Biol. Technol. Agric."},{"key":"ref_128","doi-asserted-by":"crossref","first-page":"68","DOI":"10.1049\/nbt2.12002","article-title":"Applications of Copper and Silver Nanoparticles on Wheat Plants to Induce Drought Tolerance and Increase Yield","volume":"15","author":"Ahmed","year":"2021","journal-title":"IET Nanobiotechnol."},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"100034","DOI":"10.1016\/j.plana.2023.100034","article-title":"Morpho-Physiological and Biochemical Responses in Wheat Foliar Sprayed with Zinc-Chitosan-Salicylic Acid Nanoparticles during Drought Stress","volume":"4","author":"Das","year":"2023","journal-title":"Plant Nano Biol."},{"key":"ref_130","doi-asserted-by":"crossref","first-page":"124681","DOI":"10.1016\/j.chemosphere.2019.124681","article-title":"Simultaneous Mitigation of Cadmium and Drought Stress in Wheat by Soil Application of Iron Nanoparticles","volume":"238","author":"Adrees","year":"2019","journal-title":"Chemosphere"},{"key":"ref_131","doi-asserted-by":"crossref","unstructured":"Liu, J., Chen, S.-B., Zhang, M.-C., Xiao, Y., Wang, B., Liu, H.-T., Wang, P.-F., Guo, T.-C., Kang, G.-Z., and Li, G.-Z. (2026). Application of Potassium Improves Yield and Quality Under Drought Stress by Regulating Nutrient Use Efficiency in Wheat. Plants, 15.","DOI":"10.3390\/plants15040539"},{"key":"ref_132","doi-asserted-by":"crossref","first-page":"3368","DOI":"10.1002\/agj2.21703","article-title":"Potassium Supply for Improvement of Cereals Growth Under Drought: A Review","volume":"116","author":"Damalas","year":"2024","journal-title":"Agron. J."},{"key":"ref_133","first-page":"1","article-title":"Foliar Application of Potassium and Gibberellic Acid (GA3) to Alleviate Drought Stress in Wheat","volume":"17","author":"Haque","year":"2022","journal-title":"J. Sci. Technol."},{"key":"ref_134","first-page":"175","article-title":"Potassium Applied Under Drought Improves Physiological and Nutrient Uptake Performances of Wheat (Triticum aestivum L.)","volume":"13","author":"Raza","year":"2013","journal-title":"J. Soil Sci. Plant Nutr."},{"key":"ref_135","doi-asserted-by":"crossref","first-page":"154325","DOI":"10.1016\/j.jplph.2024.154325","article-title":"Potassium May Mitigate Drought Stress by Increasing Stem Carbohydrates and Their Mobilization into Grains","volume":"303","author":"Cakmak","year":"2024","journal-title":"J. Plant Physiol."},{"key":"ref_136","doi-asserted-by":"crossref","unstructured":"Wei, J., Li, C., Li, Y., Jiang, G., Cheng, G., and Zheng, Y. (2013). Effects of External Potassium (K) Supply on Drought Tolerances of Two Contrasting Winter Wheat Cultivars. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0069737"},{"key":"ref_137","first-page":"280","article-title":"Comparative Morphological Study of Wheat (Triticum aestivum L.) Genotype\u2019s Response to Potassium Regimens Under Irrigation and Terminal Drought Conditions","volume":"7","author":"Farooq","year":"2024","journal-title":"J. Agric."},{"key":"ref_138","doi-asserted-by":"crossref","first-page":"109984","DOI":"10.1016\/j.plaphy.2025.109984","article-title":"Drought Priming Combined with Additional Potassium Application Enhances Wheat Tolerance to Post-Anthesis Drought via Osmotic Adjustment and Leaf Hydraulic Conductance","volume":"227","author":"Sun","year":"2025","journal-title":"Plant Physiol. Biochem."},{"key":"ref_139","doi-asserted-by":"crossref","unstructured":"George, N., Hany-Ali, G., Abdelhaliem, E., and Abdel-Haleem, M. (2024). Alleviating the Drought Stress and Improving the Plant Resistance Properties of Triticum aestivum via Biopriming with Aspergillus Fumigatus. BMC Plant Biol., 24.","DOI":"10.1186\/s12870-024-04840-z"},{"key":"ref_140","doi-asserted-by":"crossref","unstructured":"Al-Hawamdeh, F., Ayad, J., Alananbeh, K., and Akash, M. (2024). Bacterial Endophytes and Their Contributions to Alleviating Drought and Salinity Stresses in Wheat: A Systematic Review of Physiological Mechanisms. Agriculture, 14.","DOI":"10.3390\/agriculture14050769"},{"key":"ref_141","doi-asserted-by":"crossref","first-page":"13453","DOI":"10.15835\/nbha51413453","article-title":"The Role of Endophytes and Rhizobacteria to Combat Drought Stress in Wheat","volume":"51","author":"Mukhtiar","year":"2023","journal-title":"Not. Bot. Horti Agrobot. Cluj-Napoca"},{"key":"ref_142","doi-asserted-by":"crossref","unstructured":"Samen, F.A.E., Alsawalha, I., Alananbeh, K., Al-Karablieh, N., and Al-Abdallat, A. (2024). Evaluation of Endophytic Bacteria from Durum Wheat on Fusarium Root and Crown Rot Disease (Fusarium culmorum) Under Drought Stress. Agronomy, 14.","DOI":"10.3390\/agronomy14122912"},{"key":"ref_143","doi-asserted-by":"crossref","first-page":"44","DOI":"10.1186\/s40168-024-01770-8","article-title":"Host Genotype-Specific Rhizosphere Fungus Enhances Drought Resistance in Wheat","volume":"12","author":"Yue","year":"2024","journal-title":"Microbiome"},{"key":"ref_144","doi-asserted-by":"crossref","first-page":"012026","DOI":"10.1088\/1755-1315\/1584\/1\/012026","article-title":"Review on Synergistic Interactions of Dark Septate Endophyte in Promoting Plant Growth and Drought Resilience in the Era of Climate Change","volume":"1584","author":"Harsonowati","year":"2026","journal-title":"IOP Conf. Ser. Earth Environ. Sci."},{"key":"ref_145","doi-asserted-by":"crossref","unstructured":"Li, X., Zhang, X., Xu, M., Ye, Q., Gao, H., and He, X. (2022). Improved Tolerance of Artemisia Ordosica to Drought Stress via Dark Septate Endophyte (DSE) Symbiosis. J. Fungi, 8.","DOI":"10.3390\/jof8070730"},{"key":"ref_146","doi-asserted-by":"crossref","first-page":"e70051","DOI":"10.1111\/jac.70051","article-title":"Screening and Microcapsule Preparation of Dark Septate Endophytes for Increasing Wheat Yield Under Drought Conditions","volume":"211","author":"Ye","year":"2025","journal-title":"J. Agron. Crop Sci."},{"key":"ref_147","doi-asserted-by":"crossref","first-page":"93","DOI":"10.21608\/bfszu.2023.199664.1263","article-title":"Drought Stress Impacts and the Role of Endophytic Fungi Combating Abiotic Stress on Wheat","volume":"2023","author":"George","year":"2024","journal-title":"Bull. Fac. Sci. Zagazig Univ."},{"key":"ref_148","doi-asserted-by":"crossref","first-page":"1339","DOI":"10.1360\/SSV-2021-0313","article-title":"Effects of Five Dark Septate Endophytes Isolated from Deserts on Growing Wheat Under Drought Stress","volume":"51","author":"Liu","year":"2021","journal-title":"Sci. Sin. Vitae"},{"key":"ref_149","doi-asserted-by":"crossref","unstructured":"Yao, Z., Chen, Y., Wang, G., Hong, Y., Jiang, S., Jiang, X., Zhao, F., Zhou, C., Zhou, Y., and Tang, H. (2026). Ascomycetous Endophytic Fungi Drive Root Fungal Community Assembly in Wheat Under Moderate Drought. J. Fungi, 12.","DOI":"10.3390\/jof12020082"},{"key":"ref_150","doi-asserted-by":"crossref","first-page":"262","DOI":"10.1094\/PBIOMES-08-23-0083-R","article-title":"Multigenerational Drought Reveals a Stable Wheat Seed Fungal Community","volume":"8","author":"Becker","year":"2024","journal-title":"Phytobiomes J."},{"key":"ref_151","doi-asserted-by":"crossref","first-page":"373","DOI":"10.1007\/s11104-019-04028-7","article-title":"Alleviation of the Detrimental Effect of Water Deficit on Wheat (Triticum aestivum L.) Growth by an Indole Acetic Acid-Producing Endophytic Fungus","volume":"439","author":"Qiang","year":"2019","journal-title":"Plant Soil"},{"key":"ref_152","doi-asserted-by":"crossref","unstructured":"Wang, D., Xie, Y., Zhang, W., Yao, L., He, C., and He, X. (2024). Study on the Biological Characteristics of Dark Septate Endophytes Under Drought and Cadmium Stress and Their Effects on Regulating the Stress Resistance of Astragalus Membranaceus. J. Fungi, 10.","DOI":"10.3390\/jof10070491"},{"key":"ref_153","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1007\/s11104-019-04057-2","article-title":"Dark Septate Endophytes Improve the Growth of Host and Non-Host Plants Under Drought Stress Through Altered Root Development","volume":"439","author":"Li","year":"2019","journal-title":"Plant Soil"}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/17\/2653\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,4]],"date-time":"2026-09-04T04:29:16Z","timestamp":1788496156000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/17\/2653"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,29]]},"references-count":153,"journal-issue":{"issue":"17","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["plants15172653"],"URL":"https:\/\/doi.org\/10.3390\/plants15172653","relation":{},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,29]]}}}