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Plant Science Today (PST)

Review Articles

Vol. 13 No. sp5 (2026): Recent Advances in Agriculture

Responses of tomato to concurrent abiotic stresses, high temperature and drought: A review

DOI
https://doi.org/10.14719/pst.15617
Submitted
19 May 2026
Published
29-09-2026

Abstract

Tomato production is increasingly threatened by the growing occurrence and severity of combined drought and high-temperature stress. Although the individual effects of high temperature and drought stress on tomato growth, physiology, and productivity have been widely investigated, their combined occurrence exerts more severe and complex adverse effects on plant performance and yield. Therefore, a comprehensive understanding of their interactive impacts is essential for developing effective management strategies and breeding climate-resilient tomato cultivars. Here, we review the current understanding of the physiological, biochemical and molecular adaptive responses of tomato to combined drought and heat stresses. Co-occurring stresses during critical growth stages significantly impair photosynthesis through stomatal closure, reduced mesophyll conductance, chlorophyll degradation and disruption of photosystem II efficiency. Combined stress elevates reactive oxygen species (ROS), resulting in oxidative damage to cellular macromolecules despite the activation of antioxidant defences. Reproductive processes, such as pollen viability, fertilisation and fruit set, are highly vulnerable to combined stress. At the molecular level, stress-responsive genes and antioxidant enzymes are differentially regulated, indicating complex interactions among drought and heat signalling pathways. In addition to crop management strategies that alleviate the adverse effects of stresses to some extent, crop improvement strategies such as breeding for stress-tolerant cultivars, integrating omics insights and applying gene-editing tools offer promising avenues for enhancing tolerance to combined stresses. Deciphering integrative mechanisms governing combined drought-heat responses is pivotal for designing climate-adaptive tomato cultivars and sustaining production under global climate change.

References

  1. 1. Delarue M, Benhamed M, Fragkostefanakis S. The role of epigenetics in tomato stress adaptation. New Crops. 2025;2:100044. https://doi.org/10.1016/j.ncrops.2024.100044
  2. 2. Hoshikawa K, Pham D, Ezura H, Schafleitner R, Nakashima K. Genetic and molecular mechanisms conferring heat stress tolerance in tomato plants. Front Plant Sci. 2021;12:786688. https://doi.org/10.3389/fpls.2021.786688
  3. 3. Intergovernmental panel on climate change. 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. Lee H, Romero J, editors. Geneva (Switzerland): Intergovernmental panel on climate change; 2023. 184 p. https://doi.org/10.59327/IPCC/AR6-9789291691647
  4. 4. Mittler R. Abiotic stress, the field environment and stress combination. Trends Plant Sci. 2006;11(1):15–9. https://doi.org/10.1016/j.tplants.2005.11.002
  5. 5. Collins EJ, Bowyer C, Tsouza A, Chopra M. Tomatoes: An extensive review of the associated health impacts of tomatoes and factors that can affect their cultivation. Biology. 2022;11(2):239. https://doi.org/10.3390/biology11020239
  6. 6. Haddidi I, Duc NH, Tonk S, Rápó E, Posta K. Defense enzymes in mycorrhizal tomato plants exposed to combined drought and heat stresses. Agronomy. 2020;10(11):1657. https://doi.org/10.3390/agronomy10111657
  7. 7. Zeng ZL, Wang XQ, Zhang SB, Huang W. Mesophyll conductance limits photosynthesis in fluctuating light under combined drought and heat stresses. Plant Physiol. 2024;194(3):1498–511. https://doi.org/10.1093/plphys/kiad605
  8. 8. Pressman E, Peet MM, Pharr DM. The effect of heat stress on tomato pollen characteristics is associated with changes in carbohydrate concentration in the developing anthers. Ann Bot. 2002;90(5):631–6. https://doi.org/10.1093/aob/mcf240
  9. 9. Sato S, Kamiyama M, Iwata T, Makita N, Furukawa H, Ikeda H. Moderate increase of mean daily temperature adversely affects fruit set of Lycopersicon esculentum by disrupting specific physiological processes in male reproductive development. Ann Bot. 2006;97(5):731–8. https://doi.org/10.1093/aob/mcl037
  10. 10. Aldubai AA, Alsadon AA, Migdadi HH, Alghamdi SS, Al-Faifi SA, Afzal M. Response of tomato (Solanum lycopersicum L.) genotypes to heat stress using morphological and expression study. Plants (Basel). 2022;11(5):615. https://doi.org/10.3390/plants11050615
  11. 11. Schwarz D, Thompson AJ, Kläring HP. Guidelines to use tomato in experiments with a controlled environment. Front Plant Sci. 2014;5:625. https://doi.org/10.3389/fpls.2014.00625
  12. 12. Cammarano D, Jamshidi S, Hoogenboom G, Ruane AC, Niyogi D, Ronga D. Processing tomato production is expected to decrease by 2050 due to the projected increase in temperature. Nat Food. 2022;3:437–44. https://doi.org/10.1038/s43016-022-00521-y
  13. 13. Canene-Adams K, Campbell JK, Zaripheh S, Jeffery EH, Erdman JW Jr. The tomato as a functional food. J Nutr. 2005;135(5):1226–30. https://doi.org/10.1093/jn/135.5.1226
  14. 14. Kissoudis C, Chowdhury R, van Heusden S, van de Wiel C, Finkers R, Visser RGF, et al. Combined biotic and abiotic stress resistance in tomato. Euphytica. 2015;202:317–32. https://doi.org/10.1007/s10681-015-1363-x
  15. 15. Mustafa M, Mwangi RW, Szalai Z, Kappel N, Csambalik L. Sustainable responses to open field tomato (Solanum lycopersicum L.) stress impacts. J Agric Food Res. 2025;101825. https://doi.org/10.1016/j.jafr.2025.101825
  16. 16. Wahid A, Gelani S, Ashraf M, Foolad MR. Heat tolerance in plants: An overview. Environ Exp Bot. 2007;61(3):199–223. https://doi.org/10.1016/j.envexpbot.2007.05.011
  17. 17. Vijayakumar A, Beena R. Impact of temperature difference on the physicochemical properties and yield of tomato: A review. Chem Sci Rev Lett. 2020;9(35):665–81. https://doi.org/10.37273/chesci.CS205107159
  18. 18. Scafetta N. Impacts and risks of “realistic” global warming projections for the 21st century. Geosci Front. 2024;15(2):101774. https://doi.org/10.1016/j.gsf.2023.101774
  19. 19. Ayenan MAT, Danquah A, Hanson P, Ampomah-Dwamena C, Sodedji FAK, Asante IK, et al. Accelerating breeding for heat tolerance in tomato (Solanum lycopersicum L.): An integrated approach. Agronomy. 2019;9(11):720. https://doi.org/10.3390/agronomy9110720
  20. 20. Giordano M, Petropoulos SA, Rouphael Y. Response and defence mechanisms of vegetable crops against drought, heat and salinity stress. Agriculture. 2021;11(5):463. https://doi.org/10.3390/agriculture11050463
  21. 21. Dasgan HY, Dere S, Akhoundnejad Y, Arpaci BB. Effects of high-temperature stress during plant cultivation on tomato (Solanum lycopersicum L.) fruit nutrient content. J Food Qual. 2021;2021:7994417. https://doi.org/10.1155/2021/7994417
  22. 22. Shaheen MR, Ayyub CM, Amjad M, Waraich EA. Morpho-physiological evaluation of tomato genotypes under high temperature stress conditions. J Sci Food Agric. 2016;96(8):2698–704. https://doi.org/10.1002/jsfa.7388
  23. 23. Karkute SG, Ansari WA, Singh AK, Singh PM, Rai N, Bahadur A, et al. Characterization of high-temperature stress-tolerant tomato (Solanum lycopersicum L.) genotypes by biochemical analysis and expression profiling of heat-responsive genes. 3 Biotech. 2021;11:45. https://doi.org/10.1007/s13205-020-02587-6
  24. 24. Bhattarai S, Harvey JT, Djidonou D, Leskovar DI. Exploring morpho-physiological variation for heat stress tolerance in tomato. Plants (Basel). 2021;10(2):347. https://doi.org/10.3390/plants10020347
  25. 25. Alsamir M, Mahmood T, Trethowan R, Ahmad N. An overview of heat stress in tomato (Solanum lycopersicum L.). Saudi J Biol Sci. 2021;28(3):1654–63. https://doi.org/10.1016/j.sjbs.2020.11.088
  26. 26. Camejo D, Rodríguez P, Morales MA, Dell'Amico JM, Torrecillas A, Alarcón JJ. High temperature effects on photosynthetic activity of two tomato cultivars with different heat susceptibility. J Plant Physiol. 2005;162(3):281–9. https://doi.org/10.1016/j.jplph.2004.07.014
  27. 27. Moore CE, Meacham-Hensold K, Lemonnier P, Slattery RA, Benjamin C, Bernacchi CJ, et al. The effect of increasing temperature on crop photosynthesis: From enzymes to ecosystems. J Exp Bot. 2021;72(8):2822–44. https://doi.org/10.1093/jxb/erab090
  28. 28. Bukhov NG, Wiese C, Neimanis S, Heber U. Heat sensitivity of chloroplasts and leaves: Leakage of protons from thylakoids and reversible activation of cyclic electron transport. Photosynth Res. 1999;59:81–93. https://doi.org/10.1023/A:1006149317411
  29. 29. Nankishore A, Farrell AD. The response of contrasting tomato genotypes to combined heat and drought stress. J Plant Physiol. 2016;202:75–82. https://doi.org/10.1016/j.jplph.2016.07.006
  30. 30. Wang W, Vinocur B, Shoseyov O, Altman A. Role of plant heat-shock proteins and molecular chaperones in the abiotic stress response. Trends Plant Sci. 2004;9(5):244–52. https://doi.org/10.1016/j.tplants.2004.03.006
  31. 31. Frank G, Pressman E, Ophir R, Althan L, Shaked R, Freedman M, et al. Transcriptional profiling of maturing tomato (Solanum lycopersicum L.) microspores reveals the involvement of heat shock proteins, ROS scavengers, hormones, and sugars in the heat stress response. J Exp Bot. 2009;60(13):3891–908. https://doi.org/10.1093/jxb/erp234
  32. 32. Li H, Liu Y, Li Y, Yang Q, Yang T, Zhou Z, et al. Heat shock transcription factors regulate thermotolerance gene networks in tomato (Solanum lycopersicum) flower buds. Hortic Plant J. 2025;11(1):199–210. https://doi.org/10.1016/j.hpj.2023.06.004
  33. 33. Liang Y, Ma F, Huang S, Ma C, Shi H, Meng F, et al. Multifunctional NAC transcription factor SlJA2L integrates the ethylene pathway to orchestrate thermotolerance and fruit ripening in tomato. New Phytol. 2025;248(5):2409–27. https://doi.org/10.1111/nph.70571
  34. 34. Graci S, Barone A. Tomato plant response to heat stress: A focus on candidate genes for yield-related traits. Front Plant Sci. 2024;14:1245661. https://doi.org/10.3389/fpls.2023.1245661
  35. 35. Guo M, Liu JH, Ma X, Luo DX, Gong ZH, Lu MH. The plant heat stress transcription factors (HSFs): Structure, regulation, and function in response to abiotic stresses. Front Plant Sci. 2016;7:114. https://doi.org/10.3389/fpls.2016.00114
  36. 36. Bienert GP, Møller ALB, Kristiansen KA, Schulz A, Møller IM, Schjoerring JK, et al. Specific aquaporins facilitate the diffusion of hydrogen peroxide across membranes. J Biol Chem. 2007;282:1183–92. https://doi.org/10.1074/jbc.M603761200
  37. 37. Dere S, Kusvuran S, Dasgan HY. Does drought increase the antioxidant nutrient capacity of tomatoes? Int J Food Sci Technol. 2022;57(10):6633–45. https://doi.org/10.1111/ijfs.16008
  38. 38. Graves M, Graves T, Yuan B, Leng P. ABA-mediated reprogramming of carotenoid metabolism under heat stress impairs tomato fruit ripening. Plant Sci. 2026;362:112755.. https://doi.org/10.1016/j.plantsci.2025.112755
  39. 39. Sangu E, Tibazarwa FI, Nyomora A, Symonds RC. Expression of genes for the biosynthesis of compatible solutes during pollen development under heat stress in tomato (Solanum lycopersicum). J Plant Physiol. 2015;178:10–16. https://doi.org/10.1016/j.jplph.2015.02.002
  40. 40. Khan MA, Asaf S, Khan AL, Jan R, Kang S-M, Kim K-M, et al. Extending thermotolerance to tomato seedlings by inoculation with SA1 isolate of Bacillus cereus and comparison with exogenous humic acid application. PLoS OneXC CV. 2020;15(4):e0232228. https://doi.org/10.1371/journal.pone.0232228
  41. 41. Conti V, Parrotta L, Romi M, Del Duca S, Cai G. Tomato biodiversity and drought tolerance: A multilevel review. Int J Mol Sci. 2023;24(12):10044. https://doi.org/10.3390/ijms241210044
  42. 42. Kamanga R, Mbega E, Ndakidemi P. Drought tolerance mechanisms in plants: Physiological responses associated with water deficit stress in Solanum lycopersicum. Adv Crop Sci Tech. 2018;6(3):362. https://doi.org/10.4172/2329-8863.1000362
  43. 43. 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 (Basel). 2021;10(2):259. https://doi.org/10.3390/plants10020259
  44. 44. Hao S, Cao H, Wang H, Pan X. The physiological responses of tomato to water stress and re-water in different growth periods. Sci Hortic. 2019;249:143–54. https://doi.org/10.1016/j.scienta.2019.01.045
  45. 45. Khan TA, Saleem M, Fariduddin Q. Differential drought stress tolerance in five varieties of tomato (Solanum lycopersicum L.): An evaluation of photosynthesis and antioxidant system. J Soil Sci Plant Nutr. 2023;23:2810–31. https://doi.org/10.1007/s42729-023-01236-0
  46. 46. Liang G, Liu J, Zhang J, Guo J. Effects of drought stress on photosynthetic and physiological parameters of tomato. J Am Soc Hortic Sci. 2020;145:12–7. https://doi.org/10.21273/JASHS04725-19
  47. 47. Lamin-Samu AT, Farghal M, Ali M, Lu G. Morpho-physiological and transcriptome changes in tomato anthers of different developmental stages under drought stress. Cells. 2021;10(7):1809. https://doi.org/10.3390/cells10071809
  48. 48. Teker Yıldız M, Akı C. Evaluation of physiological and biochemical responses of four tomato (Solanum lycopersicum L.) cultivars at different drought stress levels. Agronomy. 2025;15(3):653. https://doi.org/10.3390/agronomy15030653
  49. 49. Medyouni I, Zouaoui R, Rubio E, Serino S, Ahmed HB, Bertin N. Effects of water deficit on leaves and fruit quality during the development period in tomato plant. Food Sci Nutr. 2021;9(4):1949–60. https://doi.org/10.1002/fsn3.2160
  50. 50. Niyazova NN, Huseynova IM. The antioxidant defense system of tomato (Solanum lycopersicum L.) varieties under drought stress and upon post-drought rewatering. Biochemistry (Mosc). 2024;89:1146–57. https://doi.org/10.1134/S0006297924060130
  51. 51. Xie G, Xu R, Chong L, Zhu Y. Understanding drought stress response mechanisms in tomato. Veg Res. 2024;4:33. https://doi.org/10.48130/vegres-0023-0033
  52. 52. Nicolas P, Shinozaki Y, Powell A, Philippe G, Snyder SI, Bao K, et al. Spatiotemporal dynamics of the tomato fruit transcriptome under prolonged water stress. Plant Physiol. 2022;190(4):2557–78. https://doi.org/10.1093/plphys/kiac445
  53. 53. Aslani L, Breniere T, Fanciullino AL, Bertin N. Evidence of the very early effects of water deficit on cell division and expansion processes in tomato ovaries. Environ Exp Bot. 2023;207:105227. https://doi.org/10.1016/j.envexpbot.2023.105227
  54. 54. Petrović I, Savić S, Gricourt J, Causse M, Jovanović Z, Stikić R. Effect of long-term drought on tomato leaves: The impact on metabolic and antioxidative response. Physiol Mol Biol Plants. 2021;27:2805–17. https://doi.org/10.1007/s12298-021-01102-2
  55. 55. Hernandez-Espinoza LH, Barrios-Masias FH. Physiological and anatomical changes in tomato roots in response to low water stress. Sci Hortic. 2020;265:109208. https://doi.org/10.1016/j.scienta.2020.109208
  56. 56. Pompeiano A, Moles TM, Mariotti L, Santaniello A, Di Baccio D, Scartazza A, et al. Tomato biodiversity reveals landrace enhanced drought-adaptive strategy. Plant Physiol Biochem. 2025;220:109495. https://doi.org/10.1016/j.plaphy.2025.109495
  57. 57. Patanè C, Siah S, Pellegrino A, Cosentino SL, Siracusa L. Fruit yield, polyphenols, and carotenoids in long shelf-life tomatoes in response to drought stress and rewatering. Agronomy. 2021;11(10):1943. https://doi.org/10.3390/agronomy11101943
  58. 58. Petrozza A, Summerer S, Melfi D, Mango T, Vurro F, Bettelli M, et al. A lycopene ε-cyclase TILLING allele enhances lycopene and carotenoid content in fruit and improves drought stress tolerance in tomato plants. Genes (Basel). 2023;14(6):1284. https://doi.org/10.3390/genes14061284
  59. 59. Ghannem A, Ben Aissa I, Majdoub R. Effects of regulated deficit irrigation applied at different growth stages of greenhouse grown tomato on substrate moisture, yield, fruit quality, and physiological traits. Environ Sci Pollut Res Int. 2021;28:46553–64. https://doi.org/10.1007/s11356-020-10407-w
  60. 60. Rocha Júnior DS, Barbosa ACO, Batista IA, Camillo LR, Lopes NS, Costa MGC. Impact of moderate water deficit at the fruit development stage of tomato (Solanum lycopersicum L.): Effects on plant growth, physiology, fruit yield and quality and expression of carotenoid biosynthesis genes. Acta Physiol Plant. 2023;45:65. https://doi.org/10.1007/s11738-023-03549-0
  61. 61. Liu H, Song S, Zhang H, Li Y, Niu L, Zhang J, et al. Signaling transduction of ABA, ROS, and Ca²⁺ in plant stomatal closure in response to drought. Int J Mol Sci. 2022;23(23):14824. https://doi.org/10.3390/ijms232314824
  62. 62. Shohat H, Illouz-Eliaz N, Kanno Y, Seo M, Weiss D. The tomato DELLA protein PROCERA promotes abscisic acid responses in guard cells by upregulating an abscisic acid transporter. Plant Physiol. 2020;184(1):518–28. https://doi.org/10.1104/pp.20.00485
  63. 63. Dittrich M, Mueller HM, Bauer H, Peirats-Llobet M, Rodriguez PL, Geilfus CM, et al. The role of Arabidopsis ABA receptors from the PYR/PYL/RCAR family in stomatal acclimation and closure signal integration. Nat Plants. 2019;5:1002–11. https://doi.org/10.1038/s41477-019-0490-0
  64. 64. Hsu PK, Dubeaux G, Takahashi Y, Schroeder JI. Signaling mechanisms in abscisic acid-mediated stomatal closure. Plant J. 2021;105(2):307–21. https://doi.org/10.1111/tpj.15067
  65. 65. Geiger D, Maierhofer T, Al-Rasheid KA, Scherzer S, Mumm P, Liese A, et al. Stomatal closure by fast abscisic acid signaling is mediated by the guard cell anion channel SLAH3 and the receptor RCAR1. Sci Signal. 2011;4(173):ra32 https://doi.org/10.1126/scisignal.2001346
  66. 66. Mo F, Wang M, Gao R, Gu T, Zheng K, Wang A, et al. Functional analysis of open stomata 1-slow anion channel associated 1-6 protein module in enhancing drought tolerance in tomato through stomatal regulation mechanisms. Int J Biol Macromol. 2025;308 (Pt-3):142591. https://doi.org/10.1016/j.ijbiomac.2025.142591
  67. 67. Raja V, Qadir SU, Alyemeni MN, Ahmad P. Impact of drought and heat stress individually and in combination on physio-biochemical parameters, antioxidant responses, and gene expression in Solanum lycopersicum. 3 Biotech. 2020;10:208. https://doi.org/10.1007/s13205-020-02206-4
  68. 68. Postiglione AE, Muday GK. The role of ROS homeostasis in ABA-induced guard cell signaling. Front Plant Sci. 2020;11:968. https://doi.org/10.3389/fpls.2020.00968
  69. 69. Rodrigues O, Reshetnyak G, Grondin A, Saijo Y, Leonhardt N, Maurel C, et al. Aquaporins facilitate hydrogen peroxide entry into guard cells to mediate ABA- and pathogen-triggered stomatal closure. Proc Natl Acad Sci U S A. 2017;114(34):9200–5. https://doi.org/10.1073/pnas.1704754114
  70. 70. Chen H, Shi Y, An L, Yang X, Liu J, Dai Z, et al. Overexpression of SlWRKY6 enhances drought tolerance by strengthening antioxidant defense and stomatal closure via ABA signaling in Solanum lycopersicum L. Plant Physiol Biochem. 2024;213:108855. https://doi.org/10.1016/j.plaphy.2024.108855
  71. 71. Zhao W, Huang H, Wang J, Wang X, Xu B, Yao X, et al. Jasmonic acid enhances osmotic stress responses by MYC2-mediated inhibition of protein phosphatase 2C1 and response regulators 26 transcription factor in tomato. Plant J. 2023;113(3):546–61. https://doi.org/10.1111/tpj.16067
  72. 72. Liu F, Liu Q, Wu JH, Wang ZQ, Geng YJ, Li J, et al. Arabidopsis calcineurin B-like-interacting protein kinase 8 and its functional homolog in tomato negatively regulates ABA-mediated stomatal movement and drought tolerance. Plant Cell Environ. 2024;47(7):2394–407. https://doi.org/10.1111/pce.14887
  73. 73. Khan N. Molecular insights into ABA-mediated regulation of stress tolerance and development in plants. Int J Mol Sci. 2025;26(16):7872. https://doi.org/10.3390/ijms26167872
  74. 74. Rowe JH, Topping JF, Liu J, Lindsey K. Abscisic acid regulates root growth under osmotic stress conditions via an interacting hormonal network with cytokinin, ethylene and auxin. New Phytol. 2016;211(1):225–39. https://doi.org/10.1111/nph.13882
  75. 75. Hussain Q, Asim M, Zhang R, Khan R, Farooq S, Wu J. Transcription factors interact with ABA through gene expression and signaling pathways to mitigate drought and salinity stress. Biomolecules. 2021;11(8):1159. https://doi.org/10.3390/biom11081159
  76. 76. Seo PJ, Xiang F, Qiao M, Park JY, Lee YN, Kim SG, et al. The MYB96 transcription factor mediates abscisic acid signaling during drought stress response in Arabidopsis. Plant Physiol. 2009;151(1):275–89. https://doi.org/10.1104/pp.109.144220
  77. 77. Vitale L, Francesca S, Arena C, D'Agostino N, Principio L, Vitale E, et al. Multitraits evaluation of a Solanum pennellii introgression tomato line challenged by combined abiotic stress. Plant Biol. 2023;25(4):518–28. https://doi.org/10.1111/plb.13518
  78. 78. Zhou R, Yu X, Ottosen CO, Rosenqvist E, Zhao L, Wang Y, et al. Drought stress had a predominant effect over heat stress on three tomato cultivars subjected to combined stress. BMC Plant Biol. 2017;17:24. https://doi.org/10.1186/s12870-017-0974-x
  79. 79. Heino M, Kinnunen P, Anderson W, Ray DK, Puma MJ, Varis O, et al. Increased probability of hot and dry weather extremes during the growing season threatens global crop yields. Sci Rep. 2023;13:3583. https://doi.org/10.1038/s41598-023-29378-2
  80. 80. Francesca S, Najai S, Zhou R, Decros G, Cassan C, Delmas F, et al. Phenotyping to dissect the biostimulant action of a protein hydrolysate in tomato plants under combined abiotic stress. Plant Physiol Biochem. 2022;179:32–43. https://doi.org/10.1016/j.plaphy.2022.03.012
  81. 81. Feller U. Drought stress and carbon assimilation in a warming climate: Reversible and irreversible impacts. J Plant Physiol. 2016;203:84–94. https://doi.org/10.1016/j.jplph.2016.04.002
  82. 82. Fonseca R, Micol-Ponce R, Ozuna CV, Castañeda L, Capel C, Fernández-Lozano A, et al. Resilient response to combined heat and drought stress conditions of a tomato germplasm collection, including natural and ethyl methanesulfonate-induced variants. Horticulturae. 2024;10(6):552. https://doi.org/10.3390/horticulturae10060552
  83. 83. Feller U, Vaseva II. Extreme climatic events: Impacts of drought and high temperature on physiological processes in agronomically important plants. Front Environ Sci. 2014;2:39. https://doi.org/10.3389/fenvs.2014.00039
  84. 84. Sato H, Mizoi J, Shinozaki K, Yamaguchi-Shinozaki K. Complex plant responses to drought and heat stress under climate change. Plant J. 2024;117(6):1873–92. https://doi.org/10.1111/tpj.16612
  85. 85. Bjerring Jensen N, Vrobel O, Akula Nageshbabu N, De Diego N, Tarkowski P, Ottosen CO, et al. Stomatal effects and ABA metabolism mediate differential regulation of leaf and flower cooling in tomato cultivars exposed to heat and drought stress. J Exp Bot. 2024;75(7):2156–75. https://doi.org/10.1093/jxb/erad498
  86. 86. Zhou R, Kong L, Yu X, Ottosen CO, Zhao T, Jiang F, et al. Oxidative damage and antioxidant mechanism in tomatoes responding to drought and heat stress. Acta Physiol Plant. 2019;41:20. https://doi.org/10.1007/s11738-019-2805-1
  87. 87. Wan Y, Xiao Q, Xiao X, Huang Y, Liu S, Feng W, et al. Response of tomatoes to inactivated endophyte LSE01 under combined stress of high-temperature and drought. Plant Physiol Biochem. 2024;207:108321. https://doi.org/10.1016/j.plaphy.2023.108321
  88. 88. Mumithrakamatchi AK, Alagarswamy S, Anitha K, Djanaguiraman M, Kalarani MK, Swarnapriya R, et al. Melatonin imparts tolerance to combined drought and high-temperature stresses in tomato through osmotic adjustment and ABA accumulation. Front Plant Sci. 2024;15:1382914. https://doi.org/10.3389/fpls.2024.1382914
  89. 89. Zhou R, Yu X, Wen J, Jensen NB, Dos Santos TM, Wu Z, et al. Interactive effects of elevated CO₂ concentration and combined heat and drought stress on tomato photosynthesis. BMC Plant Biol. 2020;20:260. https://doi.org/10.1186/s12870-020-02457-6
  90. 90. Yin Y, Qin K, Song X, Zhang Q, Zhou Y, Xia X, et al. BZR1 transcription factor regulates heat stress tolerance through FERONIA receptor-like kinase-mediated reactive oxygen species signaling in tomato. Plant Cell Physiol. 2018;59(11):2239–54. https://doi.org/10.1093/pcp/pcy146
  91. 91. Zhou J, Wang J, Li X, Xia XJ, Zhou YH, Shi K, et al. H2O2 mediates the crosstalk of brassinosteroid and abscisic acid in tomato responses to heat and oxidative stresses. J Exp Bot. 2014;65(15):4371–83. https://doi.org/10.1093/jxb/eru217
  92. 92. Sun M, Jiang F, Zhou R, Wen J, Cui S, Wang W, et al. Respiratory burst oxidase homologue-dependent H2O2 is essential during heat stress memory in heat sensitive tomato. Sci Hortic. 2019;258:108777. https://doi.org/10.1016/j.scienta.2019.108777
  93. 93. Flexas J, Bota J, Loreto F, Cornic G, Sharkey TD. Diffusive and metabolic limitations to photosynthesis under drought and salinity in C3 plants. Plant Biol. 2004;6(3):269–79. https://doi.org/10.1055/s-2004-820867
  94. 94. Annadurai MKK, Alagarsamy S, Karuppasami KM, Ramakrishnan S, Subramanian M, Venugopal PRB, et al. Melatonin decreases negative effects of combined drought and high temperature stresses through enhanced antioxidant defense system in tomato leaves. Horticulturae. 2023;9(6):673. https://doi.org/10.3390/horticulturae9060673
  95. 95. Mushtaq N, Wang Y, Fan J, Li Y, Ding J. Down-regulation of cytokinin receptor gene SlHK2 improves plant tolerance to drought, heat, and combined stresses in tomato. Plants (Basel). 2022;11(2):154. https://doi.org/10.3390/plants11020154
  96. 96. Duc NH, Csintalan Z, Posta K. Arbuscular mycorrhizal fungi mitigate negative effects of combined drought and heat stress on tomato plants. Plant Physiol Biochem. 2018;132:297–307. https://doi.org/10.1016/j.plaphy.2018.09.011
  97. 97. Maqsood H, Munir F, Amir R, Gul A. Genome-wide identification, comprehensive characterization of transcription factors, cis-regulatory elements, protein homology, and protein interaction network of DREB gene family in Solanum lycopersicum. Front Plant Sci. 2022;13:1031679. https://doi.org/10.3389/fpls.2022.1031679

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