Drought and high-temperature stress increasingly co-occur under climate change scenarios, posing severe threats to crop productivity globally, with particular concern for rainfed cereal systems such as wheat. The combined impact of these stresses on plant growth, development, biomass accumulation, and yield can differ substantially from their individual effects, yet evidence from diverse plant species shows that cytokinin (CK) signaling plays a pivotal role in mediating tolerance to both individual and combined stresses. Drought stress disrupts CK homeostasis by inhibiting its synthesis and accelerating its degradation, resulting in reduced CK levels in both roots and shoots. Enhancing endogenous CK levels either through exogenous application or genetic modification, such as overexpression of the isopentenyl transferase ( ipt ) gene involved in cytokinin biosynthesis, has shown promising results in improving plant stress tolerance and land-use efficiency. This review summarizes recent advances in cytokinin research related to plant stress responses and discusses prospects for its application in improving crop resilience.
Intergovernmental Panel on Climate Change (2021) Climate Change 2021—The Physical Science Basis: Working Group I Contribution to the Sixth Assessment Report of the Intergovernmental Panel on Climate Change. Cambridge University Press.
Intergovernmental Panel on Climate Change (IPCC) (2023) Climate Change 2022—Impacts, Adaptation and Vulnerability: Working Group II Contribution to the Sixth Assessment Report of the Intergovernmental Panel on Climate Change. Cambridge University Press. https://doi.org/10.1017/9781009325844
Intergovernmental Panel on Climate Change (2023) Climate Change 2023: Synthesis Report. Contribution of Working Groups I, II and III to the Sixth Assessment Report of the Intergovernmental Panel on Climate Change. Cambridge University Press.
FAO I, UNICEF, WFP and WHO (2025) The State of Food Security and Nutrition in the World 2025—Addressing High Food Price Inflation for Food Security and Nutrition. https://www.who.int/publications/m/item/the-state-of-food-security-and-nutrition-in-the-world-2025
Zhu, Y. and Gao, F. (2025) Involvement of Pathogenesis-Related Proteins and Their Roles in Abiotic Stress Responses in Plants. Biomolecules , 15, Article No. 1103. https://doi.org/10.3390/biom15081103
Qiu, R., Katul, G.G., Zhang, L., Qin, S. and Jiang, X. (2025) The Effects of Changing Environments, Abiotic Stresses, and Management Practices on Cropland Evapotranspiration: A Review. Reviews of Geophysics , 63, e2024RG000858. https://doi.org/10.1029/2024rg000858
Cao, Y., Yang, P. and Li, M. (2025) Research Progress of Peptides Discovery and Function in Resistance to Abiotic Stress in Plant. Stress Biology , 5, Article No. 36. https://doi.org/10.1007/s44154-025-00220-1
Zhao, W., Chen, X., Wang, J., Cheng, Z., Ma, X., Zheng, Q., et al . (2025) Emerging Mechanisms of Plant Responses to Abiotic Stress. Plants , 14, Article No. 3445. https://doi.org/10.3390/plants14223445
Muñoz-Espinoza, V.A., López-Climent, M.F., Casaretto, J.A. and Gómez-Cadenas, A. (2015) Water Stress Responses of Tomato Mutants Impaired in Hormone Biosynthesis Reveal Abscisic Acid, Jasmonic Acid and Salicylic Acid Interactions. Frontiers in Plant Science , 6, Article 997. https://doi.org/10.3389/fpls.2015.00997
Mittler, R. and Blumwald, E. (2010) Genetic Engineering for Modern Agriculture: Challenges and Perspectives. Annual Review of Plant Biology , 61, 443-462. https://doi.org/10.1146/annurev-arplant-042809-112116
Rizhsky, L., Liang, H., Shuman, J., Shulaev, V., Davletova, S. and Mittler, R. (2004) When Defense Pathways Collide. The Response of Arabidopsis to a Combination of Drought and Heat Stress. Plant Physiology , 134, 1683-1696. https://doi.org/10.1104/pp.103.033431
Mittler, R. (2006) Abiotic Stress, the Field Environment and Stress Combination. Trends in Plant Science , 11, 15-19. https://doi.org/10.1016/j.tplants.2005.11.002
Suzuki, N., Rivero, R.M., Shulaev, V., Blumwald, E. and Mittler, R. (2014) Abiotic and Biotic Stress Combinations. New Phytologist , 203, 32-43. https://doi.org/10.1111/nph.12797
Pradhan, G.P., Prasad, P.V.V., Fritz, A.K., Kirkham, M.B. and Gill, B.S. (2012) Effects of Drought and High Temperature Stress on Synthetic Hexaploid Wheat. Functional Plant Biology , 39, 190-198. https://doi.org/10.1071/fp11245
Prasad, K., Grigg, S.P., Barkoulas, M., Yadav, R.K., Sanchez-Perez, G.F., Pinon, V., et al . (2011) Arabidopsis PLETHORA Transcription Factors Control Phyllotaxis. Cur rent Biology , 21, 1123-1128. https://doi.org/10.1016/j.cub.2011.05.009
Wang, G.P., Zhang, X.Y., Li, F., Luo, Y. and Wang, W. (2010) Overaccumulation of Glycine Betaine Enhances Tolerance to Drought and Heat Stress in Wheat Leaves in the Protection of Photosynthesis. Photosynthetica , 48, 117-126. https://doi.org/10.1007/s11099-010-0016-5
Vile, D., Pervent, M., Belluau, M., Vasseur, F., Bresson, J., Muller, B., et al . (2011) Arabidopsis Growth under Prolonged High Temperature and Water Deficit: Independent or Interactive Effects? Plant , Cell & Environment , 35, 702-718. https://doi.org/10.1111/j.1365-3040.2011.02445.x
Rollins, J.A., Habte, E., Templer, S.E., Colby, T., Schmidt, J. and von Korff, M. (2013) Leaf Proteome Alterations in the Context of Physiological and Morphological Responses to Drought and Heat Stress in Barley ( Hordeum vulgare L.). Journal of Experimental Botany , 64, 3201-3212. https://doi.org/10.1093/jxb/ert158
Nishiyama, R., Watanabe, Y., Fujita, Y., Le, D.T., Kojima, M., Werner, T., et al . (2011) Analysis of Cytokinin Mutants and Regulation of Cytokinin Metabolic Genes Reveals Important Regulatory Roles of Cytokinins in Drought, Salt and Abscisic Acid Responses, and Abscisic Acid Biosynthesis. The Plant Cell , 23, 2169-2183. https://doi.org/10.1105/tpc.111.087395
Savchenko, G.E., Klyuchareva, E.A., Abramchik, L.M. and Serdyuchenko, E.V. (2002) Effect of Periodic Heat Shock on the Inner Membrane System of Etioplasts. Russ ian Journal of Plant Physiology , 49, 349-359. https://doi.org/10.1023/a:1015592902659
Chen, H.-H., Shen, Z.-Y. and Li, P.H. (1982) Adaptability of Crop Plants to High Temperatures Stress. Crop Science , 22, 719-725. https://doi.org/10.2135/cropsci1982.0011183x002200040006x
Martineau, J.R., Specht, J.E., Williams, J.H. and Sullivan, C.Y. (1979) Temperature Tolerance in Soybeans. I. Evaluation of a Technique for Assessing Cellular Membrane Thermostability. Crop Science , 19, 75-78. https://doi.org/10.2135/cropsci1979.0011183x001900010017x
Ashraf, M.Y., Azmi, A.R., Khan, A.H. and Ala, S. (1994) Effect of Water Stress on Total Phenols, Peroxidase Activity and Chlorophyll Content in Wheat [ Triticum aestivum L.]. Acta Physiologiae Plantarum , 16, 185-191.
Marcum, K.B. (1998) Cell Membrane Thermostability and Whole-Plant Heat Tolerance of Kentucky Bluegrass. Crop Science , 38, 1214-1218. https://doi.org/10.2135/cropsci1998.0011183x003800050017x
Ismail, A.M. and Hall, A.E. (1999) Reproductive-Stage Heat Tolerance, Leaf Membrane Thermostability and Plant Morphology in Cowpea. Crop Science , 39, 1762-1768. https://doi.org/10.2135/cropsci1999.3961762x
Wahid, A. and Shabbir, A. (2005) Induction of Heat Stress Tolerance in Barley Seedlings by Pre-Sowing Seed Treatment with Glycinebetaine. Plant Growth Regulation , 46, 133-141. https://doi.org/10.1007/s10725-005-8379-5
Grigorova, B., Vaseva, I., Demirevska, K. and Feller, U. (2011) Combined Drought and Heat Stress in Wheat: Changes in Some Heat Shock Proteins. Biologia Plantarum , 55, 105-111. https://doi.org/10.1007/s10535-011-0014-x
Machado, S. and Paulsen, G.M. (2001) Combined Effects of Drought and High Temperature on Water Relations of Wheat and Sorghum. Plant and Soil , 233, 179-187. https://doi.org/10.1023/a:1010346601643
Perdomo, J.A., Conesa, M.À., Medrano, H., Ribas-Carbó, M. and Galmés, J. (2014) Effects of Long-Term Individual and Combined Water and Temperature Stress on the Growth of Rice, Wheat and Maize: Relationship with Morphological and Physiological Acclimation. Physiologia Plantarum , 155, 149-165. https://doi.org/10.1111/ppl.12303
Rivero, R.M., Kojima, M., Gepstein, A., Sakakibara, H., Mittler, R., Gepstein, S., et al . (2007) Delayed Leaf Senescence Induces Extreme Drought Tolerance in a Flowering Plant. Proceedings of the National Academy of Sciences , 104, 19631-19636. https://doi.org/10.1073/pnas.0709453104
Wu, C., Cui, K., Wang, W., Li, Q., Fahad, S., Hu, Q., et al . (2017) Heat-Induced Cytokinin Transportation and Degradation Are Associated with Reduced Panicle Cytokinin Expression and Fewer Spikelets per Panicle in Rice. Frontiers in Plant Science , 8, Article 371. https://doi.org/10.3389/fpls.2017.00371
Werner, T., Nehnevajova, E., Köllmer, I., Novák, O., Strnad, M., Krämer, U., et al . (2010) Root-Specific Reduction of Cytokinin Causes Enhanced Root Growth, Drought Tolerance, and Leaf Mineral Enrichment in Arabidopsis and Tobacco. The Plant Cell , 22, 3905-3920. https://doi.org/10.1105/tpc.109.072694
Cortleven, A. and Schmülling, T. (2015) Regulation of Chloroplast Development and Function by Cytokinin. Journal of Experimental Botany , 66, 4999-5013. https://doi.org/10.1093/jxb/erv132
Ha, S., Vankova, R., Yamaguchi-Shinozaki, K., Shinozaki, K. and Tran, L.P. (2012) Cytokinins: Metabolism and Function in Plant Adaptation to Environmental Stresses. Trends in Plant Science , 17, 172-179. https://doi.org/10.1016/j.tplants.2011.12.005
Zwack, P.J. and Rashotte, A.M. (2015) Interactions between Cytokinin Signalling and Abiotic Stress Responses. Journal of Experimental Botany , 66, 4863-4871. https://doi.org/10.1093/jxb/erv172
Polanska, L., Vicankova, A., Novakova, M., Malbeck, J., Dobrev, P.I., Brzobohaty, B., et al . (2007) Altered Cytokinin Metabolism Affects Cytokinin, Auxin, and Abscisic Acid Contents in Leaves and Chloroplasts, and Chloroplast Ultrastructure in Transgenic Tobacco. Journal of Experimental Botany , 58, 637-649. https://doi.org/10.1093/jxb/erl235
Müller, B. and Sheen, J. (2008) Cytokinin and Auxin Interaction in Root Stem-Cell Specification during Early Embryogenesis. Nature , 453, 1094-1097. https://doi.org/10.1038/nature06943
Shashidhar, V.R., Prasad, T.G. and Sudhrshan, L. (1996) Hormone Signals from Roots to Shoots of Sunflower ( Helianthus annuus L.). Moderate Soil Drying Increases Delivery of Abscisic Acid and Depresses Delivery of Cytokinins in Xylem Sap. Annals of Botany , 78, 151-155. https://doi.org/10.1006/anbo.1996.0107
Merewitz, E.B., Du, H., Yu, W., Liu, Y., Gianfagna, T. and Huang, B. (2011) Elevated Cytokinin Content in Ipt Transgenic Creeping Bentgrass Promotes Drought Tolerance through Regulating Metabolite Accumulation. Journal of Experimental Botany , 63, 1315-1328. https://doi.org/10.1093/jxb/err372
Zhang, P., Wang, W., Zhang, G., Kaminek, M., Dobrev, P., Xu, J., et al . (2010) Senescence-Inducible Expression of Isopentenyl Transferase Extends Leaf Life, Increases Drought Stress Resistance and Alters Cytokinin Metabolism in Cassava. Journal of Integrative Plant Biology , 52, 653-669. https://doi.org/10.1111/j.1744-7909.2010.00956.x
Itai, C., Benzioni, A. and Munz, S. (1978) Heat Stress: Effects of Abscisic Acid and Kinetin on Response and Recovery of Tobacco Leaves. Plant and Cell Physiology , 19, 453-459. https://doi.org/10.1093/oxfordjournals.pcp.a075614
Rulcová, J. and Pospíšilová, J. (2001) Effect of Benzylaminopurine on Rehydration of Bean Plants after Water Stress. Biologia plantarum , 44, 75-81. https://doi.org/10.1023/a:1017922421606
Hare, P.D., Cress, W.A. and van Staden, J. (1997) The Involvement of Cytokinins in Plant Responses to Environmental Stress. Plant Growth Regulation , 23, 79-103. https://doi.org/10.1023/a:1005954525087
Brenner, E.D., Stahlberg, R., Mancuso, S., Vivanco, J., Baluška, F. and Van Volkenburgh, E. (2006) Plant Neurobiology: An Integrated View of Plant Signaling. Trends in Plant Science , 11, 413-419. https://doi.org/10.1016/j.tplants.2006.06.009
Xu, Y., Tian, J., Gianfagna, T. and Huang, B. (2009) Effects of SAG12-Ipt Expression on Cytokinin Production, Growth and Senescence of Creeping Bentgrass ( Agros tis stolonifera L.) under Heat Stress. Plant Growth Regulation , 57, 281-291. https://doi.org/10.1007/s10725-008-9346-8
Miyawaki, K., Matsumoto-Kitano, M. and Kakimoto, T. (2004) Expression of Cytokinin Biosynthetic Isopentenyltransferase Genes in Arabidopsis: Tissue Specificity and Regulation by Auxin, Cytokinin, and Nitrate. The Plant Journal , 37, 128-138. https://doi.org/10.1046/j.1365-313x.2003.01945.x
Buchanan-Wollaston, V. (1997) The Molecular Biology of Leaf Senescence. Journal of Experimental Botany , 48, 181-199. https://doi.org/10.1093/jxb/48.2.181
Van Staden, J., Cook, E.L. and Noodén, L.D. (1988) Cytokinins and Senescence. In: Noodén, L.D. and Leopold, A.C., Eds., Senescence and Aging in Plants , Elsevier, 281-328. https://doi.org/10.1016/b978-0-12-520920-5.50015-8
Gan, S. and Amasino, R.M. (1995) Inhibition of Leaf Senescence by Autoregulated Production of Cytokinin. Science , 270, 1986-1988. https://doi.org/10.1126/science.270.5244.1986
Burkhanova, E.A., Mikulovich, T.P., Kudryakova, N.V., Kukina, I.M., Smith, A.R., Hall, M.A., et al . (2001) Heat Shock Pre-Treatment Enhances the Response of Arabidopsis Thaliana Leaves and Cucurbita Pepo Cotyledons to Benzyladenine. Plant Growth Regulation , 33, 195-198. https://doi.org/10.1023/a:1017510832722
Chernyad’ev, I.I. (2005) Effect of Water Stress on the Photosynthetic Apparatus of Plants and the Protective Role of Cytokinins: A Review. Applied Biochemistry and Microbiology , 41, 115-128. https://doi.org/10.1007/s10438-005-0021-9
Skogqvist, I. (1974) Induction of Heat Sensitivity of Wheat Roots and Its Effects on Mitochondria, Adenosine Triphosphate, Triglyceride, and Total Lipid Content. E xperimental Cell Research , 86, 285-294. https://doi.org/10.1016/0014-4827(74)90715-0
Adedipe, N.O., Hunt, L.A. and Fletcher, R.A. (1971) Effects of Benzyladenine on Photosynthesis, Growth and Senescence of the Bean Plant. Physiologia Plantarum , 25, 151-153. https://doi.org/10.1111/j.1399-3054.1971.tb01105.x
Haisel, D., Pospíšilová, J., Synková, H., Schnablová, R. and Baťková, P. (2006) Effects of Abscisic Acid or Benzyladenine on Pigment Contents, Chlorophyll Fluorescence, and Chloroplast Ultrastructure during Water Stress and after Rehydration. Photos ynthetica , 44, 606-614. https://doi.org/10.1007/s11099-006-0079-5
Wahid, A., Gelani, S., Ashraf, M. and Foolad, M. (2007) Heat Tolerance in Plants: An Overview. Environmental and Experimental Botany , 61, 199-223. https://doi.org/10.1016/j.envexpbot.2007.05.011
Zavaleta-Mancera, H.A., López-Delgado, H., Loza-Tavera, H., Mora-Herrera, M., Trevilla-García, C., Vargas-Suárez, M., et al . (2007) Cytokinin Promotes Catalase and Ascorbate Peroxidase Activities and Preserves the Chloroplast Integrity during Dark-Senescence. Journal of Plant Physiology , 164, 1572-1582. https://doi.org/10.1016/j.jplph.2007.02.003
Gupta, S., Agarwal, V.P. and Gupta, N.K. (2012) Efficacy of Putrescine and Benzyladenine on Photosynthesis and Productivity in Relation to Drought Tolerance in Wheat ( Triticum aestivum L.). Physiology and Molecular Biology of Plants , 18, 331-336. https://doi.org/10.1007/s12298-012-0123-9
Shivani, N., Ajay, A., et al . (2015) Effect of Cytokinin Analogues on Cytokinin Metabolism and Stress Responsive Genes under Osmotic Stress in Wheat. The Bioscan , 10, 67-72.
Sukumar, P., Maloney, G.S. and Muday, G.K. (2013) Localized Induction of the ATP-Binding Cassette B19 Auxin Transporter Enhances Adventitious Root Formation in Arabidopsis. Plant Physiology , 162, 1392-1405. https://doi.org/10.1104/pp.113.217174
Mahan, J.R. and Mauget, S.A. (2005) Antioxidant Metabolism in Cotton Seedlings Exposed to Temperature Stress in the Field. Crop Science , 45, 2337-2345. https://doi.org/10.2135/cropsci2005.0106
Kumar, S., Kaushal, N., Nayyar, H. and Gaur, P. (2012) Abscisic Acid Induces Heat Tolerance in Chickpea ( Cicer arietinum L.) Seedlings by Facilitated Accumulation of Osmoprotectants. Acta Physiologiae Plantarum , 34, 1651-1658. https://doi.org/10.1007/s11738-012-0959-1
Zhang, X. and Ervin, E.H. (2008) Impact of Seaweed Extract-Based Cytokinins and Zeatin Riboside on Creeping Bentgrass Heat Tolerance. Crop Science , 48, 364-370. https://doi.org/10.2135/cropsci2007.05.0262
Zandalinas, S.I., Rivero, R.M., Martínez, V., Gómez-Cadenas, A. and Arbona, V. (2016) Tolerance of Citrus Plants to the Combination of High Temperatures and Drought Is Associated to the Increase in Transpiration Modulated by a Reduction in Abscisic Acid Levels. BMC Plant Biology , 16, Article No. 105. https://doi.org/10.1186/s12870-016-0791-7
Kieber, J.J. and Schaller, G.E. (2018) Cytokinin Signaling in Plant Development. Development , 145, dev149344. https://doi.org/10.1242/dev.149344
Cortleven, A., Leuendorf, J.E., Frank, M., Pezzetta, D., Bolt, S. and Schmülling, T. (2019) Cytokinin Action in Response to Abiotic and Biotic Stresses in Plants. Plant , Cell & Environment , 42, 998-1018. https://doi.org/10.1111/pce.13494
Hwang, I., Sheen, J. and Müller, B. (2012) Cytokinin Signaling Networks. Annual Review of Plant Biology , 63, 353-380. https://doi.org/10.1146/annurev-arplant-042811-105503
Devireddy, A.R., Zandalinas, S.I., Fichman, Y. and Mittler, R. (2020) Integration of Reactive Oxygen Species and Hormone Signaling during Abiotic Stress. The Plant Journal , 105, 459-476. https://doi.org/10.1111/tpj.15010
Lamaoui, M., Jemo, M., Datla, R. and Bekkaoui, F. (2018) Heat and Drought Stresses in Crops and Approaches for Their Mitigation. Frontiers in Chemistry , 6, Article 26. https://doi.org/10.3389/fchem.2018.00026
Foyer, C.H. and Noctor, G. (2020) Redox Homeostasis and Signaling in a Higher-CO 2 World. Annual Review of Plant Biology , 71, 157-182. https://doi.org/10.1146/annurev-arplant-050718-095955
Sairam, R.K. and Tyagi, A. (2004) Physiology and Molecular Biology of Salinity Stress Tolerance in Plants. Current Science , 86, 407-421.
Kishor, P.B.K., Sangam, S., Amrutha, R.N., et al . (2005) Regulation of Proline Biosynthesis, Degradation, Uptake and Transport in Higher Plants: Its Implications in Plant Growth and Abiotic Stress Tolerance. Current Science , 88, 424-438.
Santoro, M.M., Liu, Y., Khan, S.M.A., Hou, L.X. and Bolen, D.W. (1992) Increased Thermal Stability of Proteins in the Presence of Naturally Occurring Osmolytes. Bio chemistry , 31, 5278-5283. https://doi.org/10.1021/bi00138a006
Xu, J., Ji, L. and Xu, L. (2006) Lead-Induced Apoptosis in PC 12 Cells: Involvement of p53, Bcl-2 Family and Caspase-3. Toxicology Letters , 166, 160-167. https://doi.org/10.1016/j.toxlet.2006.06.643
Hasanuzzaman, M., Nahar, K., Alam, M.M. and Fujita, M. (2012) Exogenous Nitric Oxide Alleviates High Temperature Induced Oxidative Stress in Wheat ( Triticum aestivum L.) Seedlings by Modulating the Antioxidant Defense and Glyoxalase System. Australian Journal of Crop Science , 6, 1314-1323.
Almeselmani, M., Deshmukh, P. and Sairam, R. (2009) High Temperature Stress Tolerance in Wheat Genotypes: Role of Antioxidant Defence Enzymes. Acta Agronomica Hungarica , 57, 1-14. https://doi.org/10.1556/aagr.57.2009.1.1
Wang, L.J. and Li, S.H. (2006) Salicylic Acid-Induced Heat or Cold Tolerance in Relation to Ca 2+ Homeostasis and Antioxidant Systems in Young Grape Plants. Plant Science , 170, 685-694. https://doi.org/10.1016/j.plantsci.2005.09.005
Wang, Z., Pote, J. and Huang, B. (2003) Responses of Cytokinins, Antioxidant Enzymes, and Lipid Peroxidation in Shoots of Creeping Bentgrass to High Root-Zone Temperatures. Journal of the American Society for Horticultural Science , 128, 648-655. https://doi.org/10.21273/jashs.128.5.648
Synková, H., Semorádová, Š., Schnablová, R., Witters, E., Hušák, M. and Valcke, R. (2006) Cytokinin-Induced Activity of Antioxidant Enzymes in Transgenic P ssu - Ipt Tobacco during Plant Ontogeny. Biologia Plantarum , 50, 31-41. https://doi.org/10.1007/s10535-005-0071-0