Alleviation of High Temperature Stress in Bell Pepper through Foliar Application of Melatonin and Sodium Nitroprusside

Document Type : Original Article

Authors

1 Department of Agrobiotechnology, Agrarian Technological Institute, RUDN University, Moscow, Russia

2 Department of Horticulture and Landscaping, Faculty of Agriculture, University of Zabol, Zabol 98613-35856, Iran.

3 Department of Agrobiotechnology, Institute of Agriculture, RUDN University, 117198 Moscow, Russia

4 Department of Plant Protection, Faculty of Agriculture, Ferdowsi University of Mashhad, Mashhad, Iran.

Abstract
The rise in global temperatures, resulting from global warming, imposes severe stress on plants, hindering their growth and development. This study aimed to investigate the effects of melatonin and sodium nitroprusside (SNP) on the growth of California Wonder green bell pepper under heat stress conditions. A factorial experiment using a completely randomized design with three replications was conducted. Plants were exposed to temperatures of 25°C (control), 35°C, and 40°C for 24 hours following foliar application of 0 µM, 50 µM, or 100 µM melatonin and SNP. Results showed that 100 µM melatonin increased shoot dry weight by 13.26% compared to the control. Under heat stress, leaf nitrogen content increased by 32.73% and 37.24% with 50 µM and 100 µM SNP, and by 9.61% and 23.72% with 50 µM and 100 µM melatonin, respectively. At 40°C, leaf potassium levels rose significantly—up to 72% with 100 µM SNP. Additionally, 100 µM SNP increased copper and iron levels by 17.96% and 202.98%, respectively. Foliar spraying with 100 µM melatonin improved photosynthetic traits (carotenoid and carbohydrate contents) and reduced malondialdehyde levels, enhancing stress tolerance. Hydrogen peroxide content decreased by 15.16% and 20.99% with 50 µM and 100 µM SNP, respectively, at 40°C. Both melatonin and SNP significantly enhanced the activity of antioxidant enzymes (superoxide dismutase, ascorbate peroxidase, and guaiacol peroxidase) under heat stress. Overall, 100 µM melatonin was most effective in mitigating heat-induced damage and improving the physiological and biochemical performance of green bell pepper seedlings.

Keywords


  1. Abd El-Naby SKM, Esmail AMAM, Baiea MHM, et al (2020) Mitigation of Heat Stress Effects by Using Shade Net on Washington Navel Orange Trees Grown in Al-Nubaria Region, Egypt. Acta Sci. Pol. Hortorum Cultus 19:15–24. https://doi.org/10.24326/asphc.2020.03.02
  2. Afzal, S., Abdul Manap, A.S., Attiq, A., Albokhadaim, I., Kandeel, M. and Alhojaily, S.M., 2023. From imbalance to impairment: the central role of reactive oxygen species in oxidative stress-induced disorders and therapeutic exploration. Frontiers in pharmacology, 14, p.1269581.
  3. AL-Huqail AA, AL-Rashed SA, Ibrahim MM, et al (2017) Arsenic induced eco-physiological changes in Chickpea ( Cicer arietinum) and protection by gypsum, a source of sulphur and calcium. Sci Hortic 217:226–233. https://doi.org/10.1016/j.scienta.2017.02.007
  4. Altaf, M.A., Shu, H., Hao, Y., Mumtaz, M.A., Lu, X. and Wang, Z., 2022. Melatonin affects the photosynthetic performance of pepper (Capsicum annuum L.) seedlings under cold stress. Antioxidants, 11(12), p.2414.
  5. Arnao MB, Hernández-Ruiz J (2007) Melatonin promotes adventitious- and lateral root regeneration in etiolated hypocotyls of Lupinus albus L. J Pineal Res 42:147–152. https://doi.org/10.1111/j.1600-079x.2006.00396.x
  6. Astier J, Gross, I and Durner, J. (2017). Nitric oxide production in plants: an update, J Exp Bot 69:3401–3411. https://doi.org/10.1093/jxb/erx420.
  7. Awasthi R, Bhandari K, Nayyar H (2015) Temperature stress and redox homeostasis in agricultural crops. Front Environ Sci 3: https://doi.org/10.3389/fenvs.2015.00011
  8. Bajwa VS, Shukla MR, Sherif SM, et al (2014) Role of melatonin in alleviating cold stress inArabidopsis thaliana. J Pineal Res 56:238–245. https://doi.org/10.1111/jpi.12115
  9. Balfagón, D., Zandalinas, S.I., Mittler, R. and Gómez‐Cadenas, A., 2020. High temperatures modify plant responses to abiotic stress conditions. Physiologia Plantarum, 170(3), pp.335-344.
  10. Barman D, Ghimir OP, Chinnusamy V, et al (2019) Amelioration of heat stress during reproductive stage in rice by melatonin Indian J Agric Sci 89: https://doi.org/10.56093/ijas.v89i7.91688
  11. Bates LS, Waldren RP, Teare ID (1973) Rapid determination of free proline for water-stress studies. Plant Soil 39:205–207. https://doi.org/10.1007/bf00018060
  12. Beers RF, Sizer IW (1952) A spectrophotometric method for measuring the breakdown of hydrogen peroxide by catalase. , J Biol Chem.
  13. Buttar, Z.A., Wu, S.N., Arnao, M.B., Wang, C., Ullah, I. and Wang, C., 2020. Melatonin suppressed the heat stress-induced damage in wheat seedlings by modulating the antioxidant machinery. Plants, 9(7), p.809.
  14. Cakmak I, Marschner H (1992) Magnesium Deficiency and High Light Intensity Enhance Activities of Superoxide Dismutase, Ascorbate Peroxidase, and Glutathione Reductase in Bean Leaves. Plant Physiol 98:1222–1227.
  15. Campos CN, Ávila RG, de Souza KRD, et al (2019) Melatonin reduces oxidative stress and promotes drought tolerance in young Coffea arabica L. plants Agric Water Manag 211:37–47. https://doi.org/10.1016/j.agwat.2018.09.025
  16. Das K, Roychoudhury A (2014) Reactive oxygen species (ROS) and response of antioxidants as ROS-scavengers during environmental stress in plants. Front Environ Sci 2: https://doi.org/10.3389/fenvs.2014.00053
  17. Ding X, Jiang Y, He L, et al (2016) Exogenous glutathione improves high root-zone temperature tolerance by modulating photosynthesis, antioxidant and osmolytes systems in cucumber seedlings. Sci Rep6: https://doi.org/10.1038/srep35424
  18. Dong Y, Xu L, Wang Q, et al (2013) Effects of exogenous nitric oxide on photosynthesis, antioxidative ability, and mineral element contents of perennial ryegrass under copper stress J Plant Interact 9:402–411.
  19. FAO (2021) 24. FAO, 2021. Food and Agriculture Organization Corporate Statistical Database (FAOSTAT).
  20. Farooq M, Basra SMA, Wahid A, Rehman H (2009) Exogenously Applied Nitric Oxide Enhances the Drought Tolerance in Fine Grain Aromatic Rice (Oryza sativaL.). J Agron Crop Sci 195:254–261. https://doi.org/10.1111/j.1439-037x.2009.00367.x
  21. Giannopolitis CN, Ries SK (1977) Superoxide Dismutases. Plant Physiology 59:309–314. https://doi.org/10.1104/pp.59.2.309
  22. Golam Jalal Ahammed, Xu W, Liu A, Chen S (2019) Endogenous melatonin deficiency aggravates high temperature-induced oxidative stress in Solanum lycopersicum L. Environ Exp Bot 161:303–311. https://doi.org/10.1016/j.envexpbot.2018.06.006
  23. Gratani L, (1992) A non-destructive method to determine chlorophyll content of leaves. Photosynthetica 26:469–473
  24. Hakimeh Oloumi, Fatemeh Nasibi, Mozaffari H (2018) Investigation of the growth rate and secondary metabolites content of Lepidium sativum under exogenous melatonin treatment. Nova Biologica Reperta 5:144–154. https://doi.org/10.29252/nbr.5.2.144
  25. Hansika Sati, Chinchkar AV, Kataria P, Sunil Pareek (2023) Melatonin: A potential abiotic stress regulator. Plant Stress 10:100293–100293. https://doi.org/10.1016/j.stress.2023.100293
  26. Hasanuzzaman M, Nahar K, Alam Md, et al (2013) Physiological, Biochemical, and Molecular Mechanisms of Heat Stress Tolerance in Plants. Int J  Mol Sci 14:9643–9684. https://doi.org/10.3390/ijms14059643
  27. Hassan MU, Chattha MU, Khan I, et al (2020) Heat stress in cultivated plants: nature, impact, mechanisms, and mitigation strategies a review. Plant Biosystems - An International Journal Dealing with all Aspects of Plant Biology 155:211–234. https://doi.org/10.1080/11263504.2020.1727987
  28. Hong, E., Xia, X., Ji, W., Li, T., Xu, X., Chen, J., Chen, X. and Zhu, X., 2023. Effects of high temperature stress on the physiological and biochemical characteristics of Paeonia ostii. International Journal of Molecular Sciences, 24(13), p.11180.
  29. Hungria M, Kaschuk G (2014) Regulation of N2 fixation and NO3−/NH4+ assimilation in nodulated and N-fertilized Phaseolus vulgaris L. exposed to high temperature stress. Environ Exp Bot 98:32–39.
  30. Iqbal N, Fatma M, Gautam H, et al (2021) The Crosstalk of Melatonin and Hydrogen Sulfide Determines Photosynthetic Performance by Regulation of Carbohydrate Metabolism in Wheat under Heat Stress. Plants 10:1778. https://doi.org/10.3390/plants10091778
  31. Iqbal N, Sehar Z, Fatma M, et al (2022) Nitric Oxide and Abscisic Acid Mediate Heat Stress Tolerance through Regulation of Osmolytes and Antioxidants to Protect Photosynthesis and Growth in Wheat Plants. Antioxidants 11:372. https://doi.org/10.3390/antiox11020372
  32. Iqbal, N., Umar, S., Khan, N.A. and Corpas, F.J., 2021. Crosstalk between abscisic acid and nitric oxide under heat stress: exploring new vantage points. Plant Cell Reports, 40(8), pp.1429-1450.
  33. Irfan, M., El-Yazied, A.A., Sheeraz, M., Hussain, S., Sattar, A., Ali, Q., El-Gawad, H.G.A., Alzuaibr, F.M., Alharbi, M.M., Al-Balawi, S.M. and Darwish, D.B.E., 2025. Exogenous application of melatonin and jasmonic acid protects the sugar beet from heat stress by modulating the enzymatic antioxidants deference mechanism and accumulation of organic osmolytes. Acta Physiologiae Plantarum, 47(3), pp.1-15.
  34. Jamali B, Eshghi S, Kholdebarin B (2014) Response Of Strawberry “Selva” Plants On Foliar Application Of Sodium Nitroprusside (Nitric Oxide Donor) Under Saline Conditions J Hortic Res 22:139–150. https://doi.org/10.2478/johr-2014-0031
  35. Janni, M., Maestri, E., Gullì, M., Marmiroli, M. and Marmiroli, N., 2024. Plant responses to climate change, how global warming may impact on food security: A critical review. Frontiers in plant science, 14, p.1297569.
  36. Kaya A, Doganlar ZB (2019) Melatonin improves the multiple stress tolerance in pepper (Capsicum annuum). Sci Horti 256:108509. https://doi.org/10.1016/j.scienta.2019.05.036
  37. Kazemi, S., Pirmoradi, M.R., Karimi, H., Raghami, M., Rahimi, A., Kheiry, A. and Malekzadeh, M.R., 2023. Effect of foliar application of humic acid and zinc sulfate on vegetative, physiological, and biochemical characteristics of Physalis alkekengi L. under soilless culture. Journal of Soil Science and Plant Nutrition, 23(3), pp.3845-3856.
  38. Kazemi, S., Pirmoradi, M.R., Raghami, M. and Malekzadeh, M.R., 2024. Enhancing the absorption of microelements by applying humic acid and zinc sulfate in Physalis alkekengi: Improve chlorophyll content and fruit quality. Greenhouse Plant Production Journal, pp.46-82.
  39. Khan MIR, Asgher M, Khan NA (2013a) Rising temperature in the changing environment: A serious threat to plants. Climate Change and Environmental Sustainability 1:25. https://doi.org/10.5958/j.2320-6411.1.1.004
  40. Khan MIR, Iqbal N, Masood A, et al (2013b) Salicylic acid alleviates adverse effects of heat stress on photosynthesis through changes in proline production and ethylene formation. Plant Signal Behav 8:e26374. https://doi.org/10.4161/psb.26374
  41. Khan, H.M.T., Balal, R.M., Hussain, Z., Javed, S.A., Jaffar, M.T. and Alsahli, A.A., 2024. Exogenous application of melatonin mitigate the heat stress in different tomato (Solanum lycopersicum L.) cultivars. Journal of King Saud University-Science, 36(3), p.103086.
  42. Khan, H.M.T., Javed, S.A., Jaffar, M.T., Balal, R.M., ul Ain, Q., Asif, A., Shahid, M.A., El-Sheikh, M.A. and Ahmad, P., 2024. Ameliorative effect of melatonin on different tomato genotypes to induce heat stress tolerance by modulating growth and physiological attributes. Journal of King Saud University-Science, 36(10), p.103420.
  43. Kharbech O, Lamia Sakouhi, Marouane Ben Massoud, et al (2020) Nitric oxide and hydrogen sulfide protect plasma membrane integrity and mitigate chromium-induced methylglyoxal toxicity in maize seedlings. Plant Physiol Biochem 157:244–255. https://doi.org/10.1016/j.plaphy.2020.10.017
  44. Khosravi, S., Haghighi, M. and Mottaghipisheh, J., 2023. Effects of melatonin foliar application on hot pepper growth and stress tolerance. Plant Stress, 9, p.100192.
  45. Klimenko SB, Peshkova AA, Dorofeev NV (2006) NITRATE REDUCTASE ACTIVITY DURING HEAT SHOCK IN WINTER WHEAT. J stress physiol. biochem 2:50–55
  46. Kong J, Dong Y, Xu L, et al (2014) Effects of foliar application of salicylic acid and nitric oxide in alleviating iron deficiency induced chlorosis of Arachis hypogaea L. Bot Stud 55: https://doi.org/10.1186/1999-3110-55-9
  47. Korkmaz, A., Değer, Ö., Szafrańska, K., Köklü, Ş., Karaca, A., Yakupoğlu, G.Ö.K.Ç.E.N. and Kocacinar, F., 2021. Melatonin effects in enhancing chilling stress tolerance of pepper. Scientia Horticulturae, 289, p.110434.
  48. Kuppusamy, A., Alagarswamy, S., Karuppusami, K.M., Maduraimuthu, D., Natesan, S., Ramalingam, K., Muniyappan, U., Subramanian, M. and Kanagarajan, S., 2023. Melatonin enhances the photosynthesis and antioxidant enzyme activities of mung bean under drought and high-temperature stress conditions. Plants, 12(13), p.2535.
  49. Lavania D, Dhingra A, Siddiqui MH, et al (2015) Current status of the production of high temperature tolerant transgenic crops for cultivation in warmer climates. Plant Physiol Biochem 86:100–108. https://doi.org/10.1016/j.plaphy.2014.11.019
  50. Li S M, Zheng H-X, Zhan, X-S, Sui N 2021. Cytokinins as central regulators during plant growth and stress response, Plant Cell Reports. 40 : 271–282. https://doi.org/10.1007/s00299-020-02612-1.
  51. Li X, Li M-H, Deng W-W, et al (2020) Exogenous melatonin improves tea quality under moderate high temperatures by increasing epigallocatechin-3-gallate and theanine biosynthesis in Camellia sinensis L. J Plant Physiol 253:153273–153273. https://doi.org/10.1016/j.jplph.2020.153273
  52. Loreto F, Velikova V (2001) Isoprene Produced by Leaves Protects the Photosynthetic Apparatus against Ozone Damage, Quenches Ozone Products, and Reduces Lipid Peroxidation of Cellular Membranes. Plant Physiol 127:1781–1787. https://doi.org/10.1104/pp.010497
  53. Machado S, Paulsen GM (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
  54. McCready RM, Guggolz Jack, Silviera Vernon, Owens HS (1950) Determination of Starch and Amylose in Vegetables. Anal Chem 22:1156–1158. https://doi.org/10.1021/ac60045a016
  55. Mondal, S., Karmakar, S., Panda, D., Pramanik, K., Bose, B. and Singhal, R.K., 2023. Crucial plant processes under heat stress and tolerance through heat shock proteins. Plant Stress, 10, p.100227.
  56. Mostofa MG, Yoshida N, Fujita M (2013) Spermidine pretreatment enhances heat tolerance in rice seedlings through modulating antioxidative and glyoxalase systems. Plant Growth Regul 73:31–44. https://doi.org/10.1007/s10725-013-9865-9
  57. Naaz, S., Pande, A. and Laxmi, A., 2025. Nitric oxide-mediated thermomemory: a new perspective on plant heat stress resilience. Frontiers in Plant Science, 16, p.1525336.
  58. Narayanan S, Tamura PJ, Roth MR, et al (2016) Wheat leaf lipids during heat stress: I. High day and night temperatures result in major lipid alterations. Plant, Plant Cell Environ 39:787–803. https://doi.org/10.1111/pce.12649
  59. Palmieri MC, Lindermayr C, Bauwe H, et al (2010) Regulation of Plant Glycine Decarboxylase by S-Nitrosylation and Glutathionylation, Plant Physiol 152:1514–1528. https://doi.org/10.1104/pp.109.152579
  60. Parankusam S, Adimulam SS, Bhatnagar-Mathur P, Sharma KK (2017) Nitric Oxide (NO) in Plant Heat Stress Tolerance: Current Knowledge and Perspectives. Front Plant Sci 8: https://doi.org/10.3389/fpls.2017.01582
  61. Prasad A, Ferretti U, Sedlářová M, Pospíšil P (2016) Singlet oxygen production in Chlamydomonas reinhardtii under heat stress. Sci Rep 6: https://doi.org/10.1038/srep20094
  62. Rai, K.K., Pandey, N. and Rai, S.P., 2020. Salicylic acid and nitric oxide signaling in plant heat stress. Physiologia plantarum, 168(2), pp.241-255.
  63. Rao, M.J., Duan, M., Zhou, C., Jiao, J., Cheng, P., Yang, L., Wei, W., Shen, Q., Ji, P., Yang, Y. and Conteh, O., 2025. Antioxidant Defense System in Plants: Reactive Oxygen Species Production, Signaling, and Scavenging During Abiotic Stress-Induced Oxidative Damage. Horticulturae, 11(5), p.477.
  64. Raza A, Charagh S, García-Caparrós P, et al (2022) Melatonin-mediated temperature stress tolerance in plants. GM Crops Food 13:196–217. https://doi.org/10.1080/21645698.2022.2106111
  65. Ritchie SW, Nguyen HT, Holaday AS (1990) Leaf Water Content and Gas‐Exchange Parameters of Two Wheat Genotypes Differing in Drought Resistance. Crop Sci 30:105–111. https://doi.org/10.2135/cropsci1990.0011183x003000010025x
  66. Ryan J (2008) Soil And Plant Analysis: Laboratory Manual
  67. Saleem, A., Anwar, S., Nawaz, T., Fahad, S., Saud, S., Ur Rahman, T., Khan, M.N.R. and Nawaz, T., 2024. Securing a sustainable future: the climate change threat to agriculture, food security, and sustainable development goals. Journal of Umm Al-Qura University for Applied Sciences, pp.1-17.
  68. Sangu E, Tibazarwa FI, Nyomora A, Symonds RC (2015) Expression of genes for the biosynthesis of compatible solutes during pollen development under heat stress in tomato (Solanum lycopersicum). Journal of Plant Physiology 178:10–16. https://doi.org/10.1016/j.jplph.2015.02.002
  69. Santisree P, Bhatnagar-Mathur P, Sharma KK (2015) NO to drought-multifunctional role of nitric oxide in plant drought: Do we have all the answers? Plant Sci 239:44–55. https://doi.org/10.1016/j.plantsci.2015.07.012
  70. Santisree P, Bhatnagar-Mathur P, Sharma KK (2018) Molecular insights into the functional role of nitric oxide (NO) as a signal for plant responses in chickpea. Funct Plant Biol 45:267. https://doi.org/10.1071/fp16324
  71. Sehar Z, Gautam H, Masood A, Khan NA (2022) Ethylene- and Proline-Dependent Regulation of Antioxidant Enzymes to Mitigate Heat Stress and Boost Photosynthetic Efficacy in Wheat Plants J. Plant Growth Regul. https://doi.org/10.1007/s00344-022-10737-8
  72. Sehar Z, Mir IR, Khan S, et al (2023) Nitric Oxide and Proline Modulate Redox Homeostasis and Photosynthetic Metabolism in Wheat Plants under High Temperature Stress Acclimation. Plants 12:1256. https://doi.org/10.3390/plants12061256
  73. Shafeiee M, Ehsanzadeh P (2019) Physiological and biochemical mechanisms of salinity tolerance in several fennel genotypes: Existence of clearly-expressed genotypic variations. Ind Crops Prod 132:311–318.
  74. Shah Jahan M, Wang Y, Shu S, et al (2019) Exogenous salicylic acid increases the heat tolerance in Tomato (Solanum lycopersicum L) by enhancing photosynthesis efficiency and improving antioxidant defense system through scavenging of reactive oxygen species. Sci Horti 247:421–429. https://doi.org/10.1016/j.scienta.2018.12.047
  75. shaimaa Mohammed Elsayed, Saied Abd El-Naby, E.El-Gohary A, Saber Hendawy (2021) Mitigation of Heat Stress Effects on Chamomile and its Essential Oil Using Melatonin or Gibberellic Acid and some Agricultural Treatments. Egypt J Chem 0: https://doi.org/10.21608/ejchem.2021.80586.3993
  76. Shiferaw B, Baker DA (1996) An evaluation of drought screening techniques for Eragrostis tef.
  77. Siddiqui MH, Alamri SA, M. Y. Y. Al-Khaishany, et al (2017) Nitri oxide and calcium induced physio-biochemical changes in tomato (Solanum lycopersicum) plant under heat stress. Fresenius Environ Bull 26:1663–1672
  78. Song L, Ding W, Zhao M, et al (2006) Nitric oxide protects against oxidative stress under heat stress in the calluses from two ecotypes of reed. Plant Sci 171:449–458. https://doi.org/10.1016/j.plantsci.2006.05.002
  79. Soufi, H.R., Roosta, H.R., Stępień, P., Malekzadeh, K. and Hamidpour, M., 2023. Manipulation of light spectrum is an effective tool to regulate biochemical traits and gene expression in lettuce under different replacement methods of nutrient solution. Scientific Reports, 13(1), p.8600.
  80. Wahid A, Close TJ (2007) Expression of dehydrins under heat stress and their relationship with water relations of sugarcane leaves. Biol Plant 51:104–109. https://doi.org/10.1007/s10535-007-0021-0
  81. Walne, C.H. and Reddy, K.R., 2022. Temperature effects on the shoot and root growth, development, and biomass accumulation of corn (Zea mays L.). Agriculture, 12(4), p.443.
  82. Wang P, Sun X, Xie Y, et al (2014) Melatonin regulates proteomic changes during leaf senescence in Malus hupehensis. J Pineal Res 57:291–307. https://doi.org/10.1111/jpi.12169
  83. White RE (1977) Studies on mineral ion absorption by plants. Plant and Soil 46:195–208. https://doi.org/10.1007/bf00693126
  84. Xing X, Ding Y, Jin J, et al (2021) Physiological and Transcripts Analyses Reveal the Mechanism by Which Melatonin Alleviates Heat Stress in Chrysanthemum Seedlings. Front Plant Sci 12: https://doi.org/10.3389/fpls.2021.673236
  85. Xu J, JinRui X, MingWei Z, XingHua L (2005) Extraction and antioxidation of anthocyanin of black soybean seed coat.
  86. Yang Q, He H, Li H, et al (2011a) NOA1 Functions in a Temperature-Dependent Manner to Regulate Chlorophyll Biosynthesis and Rubisco Formation in Rice. PLOS ONE 6:e20015–e20015. https://doi.org/10.1371/journal.pone.0020015
  87. Yang W, Sun Y, Chen S, et al (2011b) The effect of exogenously applied nitric oxide on photosynthesis and antioxidant activity in heat stressed chrysanthemum. Biol Plant 55: https://doi.org/10.1007/s10535-011-0178-4
  88. Young LW, Wilen RW, Bonham-Smith PC (2004) High temperature stress of Brassica napus during flowering reduces micro- and megagametophyte fertility, induces fruit abortion, and disrupts seed production J Exp Bot 55:485–495. https://doi.org/10.1093/jxb/erh038
  89. Zacharoula Kostopoulou, Ioannis Therios, Roumeliotis E, et al (2015) Melatonin combined with ascorbic acid provides salt adaptation in Citrus aurantium L. seedlings. ,  Plant Physiol Biochem  86:155–165.
  90. Zahedi SM, Hosseini MS, Abadía J, Marjani M (2020) Melatonin foliar sprays elicit salinity stress tolerance and enhance fruit yield and quality in strawberry (Fragaria  ananassa Duch.). Plant Physiol Biochem 149:313–323.
  91. Zangani E, Hossein Rabbi Angourani, Babak Andalibi, Saeid Vaezi Rad, A. Mastinu, 2023 Sodium Nitroprusside Improves the Growth and Behavior of the Stomata of Silybum marianum L. Subjected to Different Degrees of Drought, Life. 13 : 875–875. https://doi.org/10.3390/life13040875.
  92. Zhang X, Ma M, Wu C, et al (2023) Mitigation of heat stress in wheat (Triticum aestivum L.) via regulation of physiological attributes using sodium nitroprusside and gibberellic acid. BMC Plant Biol 23: https://doi.org/10.1186/s12870-023-04321-9
  93. Zhang XW, Dong YJ, Qiu XK, et al (2012) Exogenous nitric oxide alleviates iron-deficiency chlorosis in peanut growing on calcareous soil. Plant Soil Environ 58:111–120. https://doi.org/10.17221/310/2011-pse
  94. Zhou X, Joshi S, Khare T, Patil S, Shang J and Kumar V (2021) Nitric oxide, crosstalk with stress regulators and plant abiotic stress tolerance, Plant Cell Rep 40:1395–1414. https://doi.org/10.1007/s00299-021-02705-5.
  95. Zhou Z, Guo K, Elbaz A, Yang Z (2009) Salicylic acid alleviates mercury toxicity by preventing oxidative stress in roots of Medicago sativa. Environ Exp Bot 65:27–34. https://doi.org/10.1016/j.envexpbot.2008.06.001

  • Receive Date 07 February 2025
  • Revise Date 16 February 2025
  • Accept Date 17 March 2025
  • First Publish Date 17 March 2025
  • Publish Date 01 March 2025