Plant Soil Environ., 2026, 72(9):542-555 | DOI: 10.17221/361/2026-PSE

Effects of drought priming at seedling stage on salt-tolerance physiology at flowering stage and subsequent seed setting of winter wheatOriginal Paper

Yiguo Liu1, Mengxin Li1, Luoyi Liu1, Fuye Bo2, Haoyang Li1, Lingyan Li1, Xuejie Wan1, Aiguo Wang1, Xiaoqing Guan3, Changhai Shi1
1 College of Agronomy, Qingdao Agricultural University, Qingdao, Shandong, P.R. China
2 Agricultural and Financial Service Centre, Linyi, Shandong, P.R. China
3 Haidu College of Qingdao Agricultural University, Qingdao, Shandong, P.R. China

To explore cultivation techniques for improving winter wheat yield in saline-alkali soils, this study investigated the effects of seedling-stage drought priming on salt tolerance and related physiological mechanisms during later growth stages. The experiment was arranged in a completely randomised design. Three soil moisture levels – 40% (W1, severe drought), 60% (W2, mild drought), and 80% (W3, well-watered) of field capacity – were applied from emergence to jointing, followed by normal irrigation and saline irrigation at 1‰ (S1), 3‰ (S2), or 5‰ (S3) NaCl. Compared with W1 and W3, mild priming (W2) significantly reduced flag leaf malondialdehyde and sterile spikelets while increasing chlorophyll and seed-setting under saline irrigation (P < 0.05). Treatment efficacy followed W2 > W3 > W1 and diminished with increasing salinity (S1 > S2 > S3). At S1, W2S1 increased stomatal conductance, superoxide dismutase (SOD) activity, and the contents of betaine and plant brassins, grains per spike (19.5%), 1 000-grain weight, and yield per plant (34.8%) versus W3S1. At S2, W2S2 increased transpiration rate, stomatal conductance, SOD activity, and the contents of soluble sugar, plant abscisic acid, and plant gibberellins, grains per spike, 1 000-grain weight, and yield per plant (35.0%) versus W3S2. Even at S3, W2S3 maintained higher SOD activity, and higher contents of osmotic substances and plant hormones (plant brassins, plant abscisic acid, and plant gibberellins), grains per spike, 1 000-grain weight, and yield per plant than W3S3. It can be seen that this advantage in seed-setting per plant is attributed to the coordinated regulation of antioxidant enzymes, osmotic substances, hormones, and photosynthesis, effectively alleviating salt stress damage. In conclusion, mild drought priming at the seedling stage establishes a stress memory that is still detectable at 7 days after anthesis (early grain filling), enhancing salt tolerance through coordinated regulation of antioxidant enzymes, osmotic substances, hormones, and photosynthesis, ultimately improving seed-setting. These findings offer a practical approach for stabilising wheat production in saline-alkali soils.

Keywords: saline-alkali stress; antioxidant defense; photosynthetic characteristics; endogenous hormones; grain formation; soil salinity

Received: August 3, 2026; Revised: September 21, 2026; Accepted: September 21, 2026; Prepublished online: September 29, 2026; Published: September 30, 2026  Show citation

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Liu Y, Li M, Liu L, Bo F, Li H, Li L, et al.. Effects of drought priming at seedling stage on salt-tolerance physiology at flowering stage and subsequent seed setting of winter wheat. Plant, Soil and Environment. 2026;72(9):542-555. doi: 10.17221/361/2026-PSE.
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References

  1. Aboud A.M.N., Safhi F.A., Alqudah A.M., Thabet S.G. (2025): Genetic associations unravel genomic regions/candidate genes of salt stress memory via enhancing antioxidant defense system in wheat. Molecular Genetics and Genomics, 300: 48. Go to original source...
  2. Ali M., Sheirf M., Amal M. (2023): Foliar spraying of nano-silicate and nano-zinc on wheat (Triticum aestivum L.) under salt stress conditions enhances productivity and reduces the optimal nitrogen fertilizer rate. Communications in Soil Science and Plant Analy-sis, 54: 2731-2744. Go to original source...
  3. Ashutus S., Jahidul K., Adnan A.P., Hasan J. (2024): Drought priming enhanced grain yield of wheat against salinity stress. Water Conservation Science and Engineering, 9: 42. Go to original source...
  4. Chang J., Wang Y., Chu J., Huang Y., Ling Y., Jiang X. (2025): Comparative salt stress responses in two turnip cultivars: insights into ion homeostasis, osmotic regulation, and antioxidant capacity. Horticulture, Environment, and Biotechnology, 66: 615-626. Go to original source...
  5. da Silva A.P.S., Alencar A.A.S., Sudré C.P., de Araújo M.S.B., Lobato A.K.S. (2024): Brassinosteroids: relevant evidence related to mitigation of abiotic and biotic stresses in plants. Agronomy, 14: 840. Go to original source...
  6. He J.W., Li Y., Wang Z.M. (2024): Drought priming promotes nitrogen use efficiency in wheat (Triticum aestivum L.) under drought stress during grain filling. Journal of Agronomy and Crop Science, 210: e12753. Go to original source...
  7. Ju F.Y., Pang J.L., Sun L.Y., Gu J.J., Wang Z., Wu X.Y., Ali S., Wang Y.H., Zhao W.Q., Wang S.S., Zhou Z.G., Chen B.L. (2023): Integrative transcriptomic, metabolomic and physiological analyses revealed the physiological and molecular mechanisms by which potassium regulates the salt tolerance of cotton (Gossypium hirsutum L.) roots. Industrial Crops and Products, 193: 116177. Go to original source...
  8. Lao C., Yu X., Zhan L., Xin P. (2024): Monitoring soil salinity in coastal wetlands with Sentinel-2 MSI data: combining fractional-order derivatives and stacked machine learning models. Agricultural Water Management, 306: 109147. Go to original source...
  9. Li C.R., Du X.J., Liu C.M. (2025a): Enhancing crop yields to ensure food security by optimizing photosynthesis. Journal of Genetics and Genomics, 52: 1082-1095. Go to original source...
  10. Li L., Zang F., Wu Q., Lu Y., Yu S., Ma Y., Zang D. (2025b): Transcriptome and metabolome analyses revealed different salt tolerance pathways in leaves and roots of Rosa rugosa Thunb. Scientia Horticulturae, 343: 114064. Go to original source...
  11. Li X., Zhang W., Niu D., Liu X.M. (2024): Effects of abiotic stress on chlorophyll metabolism. Plant Science, 342: 112030. Go to original source...
  12. Lou R., Li D., Li Y., Bian Z., Zhu Y. (2021): Effect of pre-anthesis drought hardening on post-anthesis physiological characteristics, yield and WUE in winter wheat. Phyton-International Journal of Experimental Botany, 90: 245-257. Go to original source...
  13. Louis N., Dhankher O.P., Puthur J.T. (2023): Seed priming can enhance and retain stress tolerance in ensuing generations by inducing epigenetic changes and trans-generational memory. Physiologia Plantarum, 175: e13881. Go to original source... Go to PubMed...
  14. Mushtap A., Ayesha, Sabir (2021): Effect of sodium silicate and salicylic acid on sodium and potassium ratio in wheat (Triticum aes-tivum L.) grown under salt stress. Silicon, 14: 5595-5600. Go to original source...
  15. Rao D.M., Dong Q.S., Cheng T., Yu D.B., Qi L.Q., Zhao J., Yan X.Y., Meng F.G., Zhang W. (2024): Research progress of crop photo-synthetic efficiency adaptation to adversity and improving ways. Plant Physiology Journal, 60: 1229-1240. (In Chinese)
  16. Ping L., Hong Y., Lu W., Wang L., Liu H.J., Huo H.Q., Zhang C.J., Liu A.Z., Zhu A.D., Hu J.Y., Lin Y.J., Liu L. (2019): Physiological and transcriptome analyses reveal short-term responses and formation of memory under drought stress in rice. Frontiers in Genetics, 10: 55. Go to original source... Go to PubMed...
  17. Rio N., Taheb M.S., Waisuddin A., Sato K., Kawaura K. (2022): Comparative transcriptome analysis of synthetic and common wheat in response to salt stress. Scientific Reports, 12: 11534. Go to original source... Go to PubMed...
  18. Ru C., Hu X.T., Wang W.N. (2024): Repeated drought stimuli increased wheat yield and drought tolerance by improving photosynthe-sis and antioxidant defense during recovery. Plant and Soil, 507: 533-556. Go to original source...
  19. Sarika K., Nazir F., Jain K., Khan M.I.R. (2023): GABA and potassium modulates defence systems, assimilation of nitrogen and carbon, and yield traits under salt stress in wheat. Journal of Plant Growth Regulation, 42: 6721-6740. Go to original source...
  20. Schat L., Schubert M., Fjellheim S., Humphreys A.M. (2025): Drought tolerance as an evolutionary precursor to frost and winter tolerance in grasses. Evolution, 79: 654-665. Go to original source... Go to PubMed...
  21. Simkin A.J., Kapoor L., Doss C.G.P., Hofmann T.A., Lawson T., Ramamoorthy S. (2022): The role of photosynthesis related pigments in light harvesting, photoprotection and enhancement of photosynthetic yield in planta. Photosynthesis Research, 152: 23-42. Go to original source... Go to PubMed...
  22. Wang X., Ma G., Feng N., Zheng D., Zhou H., Li J., Wu J., Xu B., Su W., Huang Y. (2025): Abscisic acid can improve the salt toler-ance and yield of rice by improving its physiological characteristics. Agronomy, 15: 309. Go to original source...
  23. Wang X.Y., Li X.M., Zhao W., Hou X.M., Dong S.K. (2024): Current views of drought research: experimental methods, adaptation mechanisms and regulatory strategies. Frontiers in Plant Science, 15: 1371895. Go to original source... Go to PubMed...
  24. Wang Z.H., Wei Y.Q., Zhao Y.R., Wang Y.J., Zou F., Huang S.Q., Yang X.L., Xu Z.W., Hu H. (2022): Physiological and transcriptional evaluation of sweet sorghum seedlings in response to single and combined drought and salinity stress. South African Journal of Bota-ny, 146: 459-471. Go to original source...
  25. Xiao W., Marija V., Dong J. (2014): Improved tolerance to drought stress after anthesis due to priming before anthesis in wheat (Triti-cum aestivum L.) var. Vinjett. Journal of Experimental Botany, 65: 6441-6456. Go to original source... Go to PubMed...
  26. Yong S.Y., Chen Q., Xu F., Fu H., Liang G.L., Guo Q.G. (2024): Exploring the interplay between angiosperm chlorophyll metabolism and environmental factors. Planta: An International Journal of Plant Biology, 260: 25. Go to original source... Go to PubMed...
  27. Yoshida T., Obata T., Fernie A.R. (2024): Hormonal regulation of plant primary metabolism under drought. Journal of Experimental Botany, 75: 1714-1725. Go to original source... Go to PubMed...
  28. Zhang Q.H., Huo Y.F., Dong M.Q., Li F.Y., Wang X.Y., Li L., Cai H.J. (2024a): Effects of different degrees and frequencies of drought exercise on water productivity in winter wheat. Journal of Triticeae Crops, 44: 1515-1527. (In Chinese)
  29. Zhang W.M., Zou Y.F., Zhou W.L., Li C., Zuo L.J., Miao L.J., Cui X.F. (2024b): Current status and obstacles of narrowing yield gaps of four major crops. Journal of the Science of Food and Agriculture, 105: 42-53. Go to original source... Go to PubMed...

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