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Plant Science Today (PST)

Review Articles

Vol. 13 No. 3 (2026)

Morpho-physiological and biochemical adaptation of wheat (Triticum aestivum L.) under drought stress: An integrated review

DOI
https://doi.org/10.14719/pst.12382
Submitted
22 October 2025
Published
17-09-2026 — Updated on 30-09-2026
Versions

Abstract

Drought stress is the major factor that causes yield reduction worldwide. Depending on the crop and the intensity of the drought, yield losses of 30–90 % have been reported. Wheat specifically loses 50–60 % of its yield, which may exceed up to 60 % under severe drought. Wheat triggers a wide range of changes under drought stress. Understanding the morphological, physiological and biochemical responses of wheat to drought stress is essential for developing resilient cultivars and improving crop management. This review explores key adaptations that help wheat optimise water uptake and minimise water loss under drought conditions and understanding of how these mechanisms interact as an integrated adaptive network under drought conditions. Morphological changes such as seed germination interference (20–30 %), shoot and root system adjustments have been observed and discussed in detail, followed by physiological responses such as plant-water relation effect of drought on which include reduction in photosynthesis (30–70 %), chlorophyll content (20–40 %) and enhanced osmotic adjustment and hormonal signalling are also investigated. Additionally, biochemical changes such as antioxidant production and Reactive Oxygen Species (ROS) have been explored thoroughly. Collectively, these adaptation responses, constitute a dynamic and extensive network of actions intended for the mitigation of drought-induced damages. By comprehending these mechanisms, we may better understand resilience of wheat and provide the groundwork for creating drought-tolerant cultivars using conventional and novel breeding techniques as well as biotechnological interventions. The aim to underscore the significance of connecting morpho-physiological and biochemical traits to enhance the sustainability of wheat under water-limited environments, highlighting their interconnections relevant for developing resilient cultivars.

References

  1. 1. Anjum SA, Xie XY, Wang LC, Saleem MF, Man C, Lei W. Morphological, physiological and biochemical responses of plants to drought stress. Afr J Agric Res. 2011;6. https://doi.org/10.5897/AJAR10.027
  2. 2. Duvnjak J, Lončarić A, Brkljačić L, Šamec D, Šarčević H, Salopek-Sondi B, et al. Morpho-physiological and hormonal response of winter wheat varieties to drought stress at stem elongation and anthesis stages. Plants. 2023;12:418. https://doi.org/10.3390/plants12030418
  3. 3. Sahoo SS, Sahu DP, Behra RK. Drought stress mitigation in wheat seedlings via green synthesized silver nanoparticle priming. Seeds. 2025;4(4):1-27. https://doi.org/10.3390/seeds4040062
  4. 4. Shinta, Bentoy KMY, Fauzia AN, Nampei M, Linh NM, Ueda A. Physiological and transcriptomic insights into mechanisms of salt tolerance in the leaves and roots of the halophyte Suaeda japonica Makino under high salinity stress. J Plant Growth Regul. 2026;45:321-39. https://doi.org/10.1007/s00344-025-11864-8
  5. 5. Priyanka, Rana V, Basandrai D, Sood VK, Rana A. Identification of promising and stable genotypes of wheat (Triticum aestivum L.) for grain yield under varied climatic conditions of north-western Himalayas using joint regression analysis. J Cereal Res. 2024;16(1):44-52. https://doi.org/10.25174/2582-2675/2024/136259
  6. 6. Pinto RS, Reynolds MP, Mathews KL, McIntyre CL, Olivares-Villegas JJ, Chapman SC. Heat and drought adaptive QTL in a wheat population designed to minimize confounding agronomic effects. Theor Appl Genet. 2010;121(6):1001-21. https://doi.org/10.1007/s00122-010-1351-4
  7. 7. Yadav AK, Carroll AJ, Estavillo GM, Rebetzke GJ, Pogson BJ. Wheat drought tolerance in the field is predicted by amino acid responses to glasshouse-imposed drought. J Exp Bot. 2019;70(18):4931-48. https://doi.org/10.1093/jxb/erz224
  8. 8. Page MJ, McKenzie JE, Bossuyt PM, Boutron I, Hoffmann TC, Mulrow CD, et al. The PRISMA 2020 statement: an updated guideline for reporting systematic reviews. BMJ. 2021;372:n71. https://doi.org/10.1136/bmj.n71
  9. 9. Disasa KN, Yan H, Zhang J, Wang G, Zhang C, Zhang D, et al. Comprehensive review of drought characteristics and intensification under climate change: implications for agriculture and water resources. J Hydrol. 2026;664:134571. https://doi.org/10.1016/j.jhydrol.2025.134571
  10. 10. Nyaupane S, Poudel MR, Panthi B, Dhakal A, Paudel H, Bhandari R. Drought stress effect, tolerance and management in wheat: a review. Cogent Food Agric. 2024;10. https://doi.org/10.1080/23311932.2023.2296094
  11. 11. Seleiman MF, Al-Suhaibani N, Ali N, Akmal M, Alotaibi M, Refay Y, et al. Drought stress impacts on plants and different approaches to alleviate its adverse effects. Plants. 2021;10:259. https://doi.org/10.3390/plants10020259
  12. 12. Tatar O, Brück H, Asch F. Photosynthesis and remobilization of dry matter in wheat as affected by progressive drought stress at stem elongation stage. J Agron Crop Sci. 2016;202:292-99. https://doi.org/10.1111/jac.12160
  13. 13. Poudel MR, Ghimire S, P MP, Ey, Dhakal KH, Thapa DB, et al. Evaluation of wheat genotypes under irrigated, heat stress and drought conditions. J Biol Todays World. 2020;9:1-12. https://doi.org/10.35248/2322-3308.20.9.212
  14. 14. Gašparíková O, Mistrík I. Review of Plant Roots: The Hidden Half. Ann Bot. 2002;90:775. https://doi.org/10.1093/aob/mcf252
  15. 15. Kang J, Peng Y, Xu W. Crop root responses to drought stress: molecular mechanisms, nutrient regulations and interactions with microorganisms in the rhizosphere. Int J Mol Sci. 2022;23:9310. https://doi.org/10.3390/ijms23169310
  16. 16. Ali M, Fareed A, Jan SU, Gul A, Gul S, Nadeem M, et al. Cellular mechanisms of drought tolerance in wheat. In: Climate Change and Food Security with Emphasis on Wheat. 2020. p. 155-67. https://doi.org/10.1016/B978-0-12-819527-7.00009-1
  17. 17. Henry A, Cal AJ, Batoto TC, Torres RO, Serraj R. Root attributes affecting water uptake of rice (Oryza sativa) under drought. J Exp Bot. 2012;63:4751-63. https://doi.org/10.1093/jxb/ers150
  18. 18. Ahmad Z, Waraich EA, Akhtar S, Anjum S, Ahmad T, Mahboob W, et al. Physiological responses of wheat to drought stress and its mitigation approaches. Acta Physiol Plant. 2018;40:1-13. https://doi.org/10.1007/s11738-018-2651-6
  19. 19. Nayyar H, Gupta D. Differential sensitivity of C3 and C4 plants to water deficit stress: association with oxidative stress and antioxidants. Environ Exp Bot. 2006;58:106-13. https://doi.org/10.1016/j.envexpbot.2005.06.021
  20. 20. Sourour A, Afef O, Mounir R, Younes Mongi B. A review: morphological, physiological, biochemical and molecular plant responses to water deficit stress. https://doi.org/10.9790/1813-0601010104
  21. 21. Maghsoudi K, Emam Y, Pessarakli M. Effect of silicon on photosynthetic gas exchange, photosynthetic pigments, cell membrane stability and relative water content of different wheat cultivars under drought stress conditions. J Plant Nutr. 2016;39:1001-15. https://doi.org/10.1080/01904167.2015.1109108
  22. 22. Mao H, Li S, Wang Z, Cheng X, Li F, Mei F, et al. Regulatory changes in TaSNAC8–6A are associated with drought tolerance in wheat seedlings. Plant Biotechnol J. 2020;18:1078-92. https://doi.org/10.1111/pbi.13277
  23. 23. Farooq M, Wahid A, Kobayashi N, Fujita D, Basra SMA. Plant drought stress: effects, mechanisms and management. Agron Sustain Dev. 2009;29:185-212. https://doi.org/10.1051/agro:2008021
  24. 24. Chang H, Chen D, Kam J, Richardson T, Drenth J, Guo X, et al. Abiotic stress upregulated TaZFP34 represses the expression of type-B response regulator and SHY2 genes and enhances root to shoot ratio in wheat. Plant Sci. 2016;252:88-102. https://doi.org/10.1016/j.plantsci.2016.07.011
  25. 25. Nikolaeva MK, Maevskaya SN, Shugaev AG, Bukhov NG. Effect of drought on chlorophyll content and antioxidant enzyme activities in leaves of three wheat cultivars varying in productivity. Russ J Plant Physiol. 2010;57:87-95. https://doi.org/10.1134/S1021443710010127
  26. 26. Bhanbhro N, Wang HJ, Yang H, Xu XJ, Jakhar AM, Shalmani A, et al. Revisiting the molecular mechanisms and adaptive strategies associated with drought stress tolerance in common wheat (Triticum aestivum L.). Plant Stress. 2024;11:100298. https://doi.org/10.1016/j.stress.2023.100298
  27. 27. Yu J, Jiang M, Guo C. Crop pollen development under drought: from the phenotype to the mechanism. Int J Mol Sci. 2019;20:1550. https://doi.org/10.3390/ijms20071550
  28. 28. Nezhadahmadi A, Prodhan ZH, Faruq G. Drought tolerance in wheat. Sci World J. 2013;2013:610721. https://doi.org/10.1155/2013/610721
  29. 29. Moayedi AA, Boyce AN, Barakbah SS. The performance of durum and bread wheat genotypes associated with yield and yield component under different water deficit conditions. Aust J Crop Sci. 2010;4(1):70-77.
  30. 30. Zhang Z, Hu M, Xu W, Wang Y, Huang K, Zhang C, et al. Understanding the molecular mechanism of anther development under abiotic stresses. Plant Mol Biol. 2021;105:1-10. https://doi.org/10.1007/s11103-020-01074-z
  31. 31. Mao H, Jian C, Cheng X, Chen B, Mei F, Li F, et al. The wheat ABA receptor gene TaPYL1–1B contributes to drought tolerance and grain yield by increasing water-use efficiency. Plant Biotechnol J. 2022;20:846-61. https://doi.org/10.1111/pbi.13764
  32. 32. Saed-Moucheshi A, Sohrabi F, Fasihfar E, Baniasadi F, Riasat M, Mozafari AA. Superoxide dismutase (SOD) as a selection criterion for triticale grain yield under drought stress: a comprehensive study on genomics and expression profiling, bioinformatics, heritability and phenotypic variability. BMC Plant Biol. 2021;21:1-19. https://doi.org/10.1186/s12870-021-02919-5
  33. 33. Hashmi QA, Ahmad M, Ahmad E, Tariq M, Rafeh A, Awais M. Exploring drought tolerance in wheat: insights from biochemical, morphological and physiological responses. Asian J Res Crop Sci. 2023;8:344-60. https://doi.org/10.9734/ajrcs/2023/v8i4216
  34. 34. Gao H, Wang Y, Xu P, Zhang Z. Overexpression of a WRKY transcription factor TaWRKY2 enhances drought stress tolerance in transgenic wheat. Front Plant Sci. 2018;9:997. https://doi.org/10.3389/fpls.2018.00997
  35. 35. Li H, Li L, ShangGuan G, Jia C, Deng S, Noman M, et al. Genome-wide identification and expression analysis of bZIP gene family in Carthamus tinctorius L. Sci Rep. 2020;10. https://doi.org/10.1038/s41598-020-72390-z
  36. 36. Qayyum A, Al Ayoubi S, Sher A, Bibi Y, Ahmad S, Shen Z, et al. Improvement in drought tolerance in bread wheat is related to an improvement in osmolyte production, antioxidant enzyme activities and gaseous exchange. Saudi J Biol Sci. 2021;28:5238-49. https://doi.org/10.1016/j.sjbs.2021.05.040
  37. 37. Xue GP, Way HM, Richardson T, Drenth J, Joyce PA, McIntyre CL. Overexpression of TaNAC69 leads to enhanced transcript levels of stress up-regulated genes and dehydration tolerance in bread wheat. Mol Plant. 2011;4:697-712. https://doi.org/10.1093/mp/ssr013
  38. 38. Li Y, Zhang S, Zhang N, Zhang W, Li M, Liu B, et al. MYB-CC transcription factor, TaMYBsm3, cloned from wheat is involved in drought tolerance. BMC Plant Biol. 2019;19:1-11. https://doi.org/10.1186/s12870-019-1751-9
  39. 39. Sivamani E, Bahieldin A, Wraith JM, Al-Niemi T, Dyer WE, Ho THD, et al. Improved biomass productivity and water use efficiency under water deficit conditions in transgenic wheat constitutively expressing the barley HVA1 gene. Plant Sci. 2000;155:1-9. https://doi.org/10.1016/S0168-9452(99)00247-2
  40. 40. Shiqing G, Huijun X, Xianguo C, Ming C, Zhaoshi X, Liancheng L, et al. Improvement of wheat drought and salt tolerance by expression of a stress-inducible transcription factor GmDREB of soybean (Glycine max). Chin Sci Bull. 2005;50:2714-23. https://doi.org/10.1007/BF02899641
  41. 41. Pellegrineschi A, Reynolds M, Pacheco M, Brito RM, Almeraya R, Yamaguchi-Shinozaki K, et al. Stress-induced expression in wheat of the Arabidopsis thaliana DREB1A gene delays water stress symptoms under greenhouse conditions. Genome. 2004;47:493-500. https://doi.org/10.1139/g03-140
  42. 42. Yu TF, Xu ZS, Guo JK, Wang YX, Abernathy B, Fu JD, et al. Improved drought tolerance in wheat plants overexpressing a synthetic bacterial cold shock protein gene SeCspA. Sci Rep. 2017;7:1-14. https://doi.org/10.1038/srep44050
  43. 43. Gupta PK. Drought-tolerant transgenic wheat HB4®: a hope for the future. Trends Biotechnol. 2024;42:807-09. https://doi.org/10.1016/j.tibtech.2023.12.007
  44. 44. Tian F, Wang W, Liang C, Wang X, Wang G, Wang W. Overaccumulation of glycine betaine makes the function of the thylakoid membrane better in wheat under salt stress. Crop J. 2017;5:73-82. https://doi.org/10.1016/j.cj.2016.05.008
  45. 45. Fehér-Juhász E, Majer P, Sass L, Lantos C, Csiszár J, Turóczy Z, et al. Phenotyping shows improved physiological traits and seed yield of transgenic wheat plants expressing the alfalfa aldose reductase under permanent drought stress. Acta Physiol Plant. 2014;36:663-73. https://doi.org/10.1007/S11738-013-1445-0
  46. 46. El-Esawi MA, Al-Ghamdi AA, Ali HM, Ahmad M. Overexpression of AtWRKY30 transcription factor enhances heat and drought stress tolerance in wheat (Triticum aestivum L.). Genes. 2019;10(2):163. https://doi.org/10.3390/genes10020163
  47. 47. Li L, Zheng M, Deng G, Liang J, Zhang H, Pan Z, et al. Overexpression of AtHDG11 enhanced drought tolerance in wheat (Triticum aestivum L.). Mol Breed. 2016;36:1-10. https://doi.org/10.1007/s11032-016-0447-1
  48. 48. Abdallah NA, Elsharawy H, Abulela HA, Thilmony R, Abdelhadi AA, Elarabi NI. Multiplex CRISPR/Cas9-mediated genome editing to address drought tolerance in wheat. GM Crops Food. 2025;16:1-17. https://doi.org/10.1080/21645698.2022.2120313
  49. 49. Chaouachi L, Guzmán-López MH, Karmous C, Barro F, Marín-Sanz M. Assessing drought stress response in low-gliadin wheat developed via RNAi and CRISPR/Cas. Plant Stress. 2025;16:100906. https://doi.org/10.1016/j.stress.2025.100906
  50. 50. Chen K, Wang Y, Zhang R, Zhang H, Gao C. CRISPR/Cas genome editing and precision plant breeding in agriculture. Annu Rev Plant Biol. 2019;70:667-97. https://doi.org/10.1146/annurev-arplant-050718-100049
  51. 51. Joga RJ, Zotti MJ, Smagghe G, Christiaens O. RNAi efficiency, systemic properties and novel delivery methods of pest insect control: what we know so far. Front Physiol. 2016;7:553. https://doi.org/10.3389/fphys.2016.00553
  52. 52. Lamaoui M, Jemo M, Datla R, Bekkaoui F. Heat and drought stress in crops and approaches for their mitigation. Front Chem. 2018;6:26. https://doi.org/10.3389/fchem.2018.00026

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