Journals / Anadolu Tarım Bilimleri Dergisi / 2025 / Cilt: 41

Effects of Combined Drought and Salinity Stress on Oxidative Damage and the Antioxidant Enzyme System in Aronia (Aronia melanocarpa L.) ‘Nero’ under In Vitro Conditions

In Vitro Koşullarda Kombine Kuraklık ve Tuz Stresinin Aronya (Aronia melanocarpa L.) da Oksidatif Hasar ve Antioksidan Enzim Sistemi Üzerine Etkisi

Çağlar Kaya(ÇANAKKALE ONSEKİZ MART ÜNİVERSİTESİ)0000-0002-7054-3081
Pages
—
Publication Date
2026-10-16
🏛️Bibliographic metadata and full text access for this article are provided via DergiPark.
DergiPark Record ↗
✨

AI Summary & Key Takeaways

AI Insight
⚡ TL;DR(Too Long; Didn't Read)

Future studies should evaluate different stress intensities and exposure durations across various aronia cultivars and under field conditions, while also investigating the molecular and metabolic mechanisms underlying tolerance to combined stress.

📌 3 Key Takeaways

🎯
Objective & Hypothesis:This study investigated the effects of single and combined drought (10% PEG-6000) and salinity (100 mM NaCl) stress on oxidative injury, antioxidant defense, and physiological responses of aronia (Aronia melanocarpa L.) ‘Nero’ under in-vitro conditions.
📊
Key Findings & Data:After 14 days, combined stress induced markedly stronger biochemical disruption than individual treatments.
💡
Conclusion & Impact:Future studies should evaluate different stress intensities and exposure durations across various aronia cultivars and under field conditions, while also investigating the molecular and metabolic mechanisms underlying tolerance to combined stress.
🔬Design:Araştırma Makalesi
📍Scope:Comprehensive Sample
⚙️Variables:Kombine abiyotik stres · Oksidatif hasar · Antioksidan enzim sistemi

Abstract

This study investigated the effects of single and combined drought (10% PEG-6000) and salinity (100 mM NaCl) stress on oxidative injury, antioxidant defense, and physiological responses of aronia (Aronia melanocarpa L.) ‘Nero’ under in-vitro conditions. After 14 days, combined stress induced markedly stronger biochemical disruption than individual treatments. Malondialdehyde (MDA) increased from 8.4 nmol g-1 FW in the control to 14.7 (PEG) and 16.2 (NaCl), reaching 24.8 nmol g-1 FW under combined stress. Hydrogen peroxide (H2O2) rose from 3.2 to 11.6 µmol g-1 FW in the dual treatment, indicating intensified reactive oxygen species accumulation. Antioxidant enzymes were strongly upregulated in response to stress. SOD activity increased from 62.4 to 138.6 U mg-1 protein under combined stress, while CAT (21.8 U mg-1 protein), APX (13.2 U mg-1 protein), GR (9.7 U mg-1 protein), and POD (29.4 U mg-1 protein) also showed their highest activities in the dual treatment. Non-enzymatic antioxidants followed a similar pattern: proline increased nearly fivefold (1.8 to 8.7 µmol g-1 FW), total phenolics rose from 5.4 to 13.8 mg GAE g-1 FW, flavonoids reached 5.2 mg CE g-1 FW, and DPPH scavenging activity increased from 38.6% to 64.3%. Despite enhanced antioxidant responses, oxidative markers remained elevated under combined stress. Relative water content declined from 91.4% to 61.3%, and SPAD values decreased from 42.5 to 28.9, indicating impaired water status and chlorophyll stability. Simultaneous osmotic and ionic stress, therefore, imposes a substantially greater redox and physiological burden than single stress exposure in aronia ‘Nero’. Future studies should evaluate different stress intensities and exposure durations across various aronia cultivars and under field conditions, while also investigating the molecular and metabolic mechanisms underlying tolerance to combined stress.

Özet

Bu çalışma, in vitro koşullarda aronya (Aronia melanocarpa L.) ‘Nero’ çeşidinde tekil ve kombine kuraklık (%10 PEG-6000) ile tuz (100 mM NaCl) stresinin oksidatif hasar, antioksidan savunma sistemi ve fizyolojik yanıtlar üzerindeki etkilerini araştırmıştır. On dört günlük uygulama sonunda, kombine stresin tekil uygulamalara kıyasla belirgin düzeyde daha güçlü biyokimyasal bozulmaya yol açtığı belirlenmiştir. Malondialdehit (MDA) içeriği kontrolde 8.4 nmol g-1 TA iken, PEG uygulamasında 14.7 ve NaCl uygulamasında 16.2 nmol g-1 TA’ya yükselmiş, kombine stres altında ise 24.8 nmol g-1 TA’ya ulaşmıştır. Hidrojen peroksit (H2O2) düzeyi ise 3.2 µmol g-1 TA’dan çift stres uygulamasında 11.6 µmol g-1 TA’ya yükselerek reaktif oksijen türlerinin belirgin birikimini göstermiştir. Antioksidan enzim aktiviteleri stres koşullarında güçlü biçimde artış göstermiştir. SOD aktivitesi kombine stres altın 62.4’ten 138.6 U mg-1 protein’e yükselmiştir. Benzer şekilde CAT (21.8 U mg-1 protein), APX (13.2 U mg-1 protein), GR (9.7 U mg-1 protein) ve POD (29.4 U mg-1 protein) aktiviteleri de en yüksek değerlere kombine stres uygulamasında ulaşmıştır. Enzimatik olmayan antioksidanlar da benzer bir eğilim göstermiştir. Prolin miktarı yaklaşık beş kat artarak 1.8’den 8.7 µmol g-1 TA’ya yükselmiştir. Toplam fenolik içerik 5.4’ten 13.8 mg GAE g-1 TA’ya çıkmış, flavonoid içeriği 5.2 mg CE g-1 TA’ya ulaşmış ve DPPH radikal süpürme aktivitesi %38.6’dan %64.3’e yükselmiştir. Bununla birlikte, antioksidan yanıtların artmasına rağmen oksidatif belirteçlerin kombine stres altında yüksek kalması dikkat çekmiştir. Relatif su içeriği %91.4’ten %61.3’e gerilemiş, SPAD değerleri ise 42.5’ten 28.9’a düşerek su durumu ve klorofil stabilitesinde belirgin bir bozulmaya işaret etmiştir. Elde edilen bulgular, ozmotik ve iyonik stresin eş zamanlı uygulanmasının aronya ‘Nero’ bitkisinde tekil stres uygulamalarına kıyasla daha ağır bir redoks ve fizyolojik yük oluşturduğunu göstermektedir. Gelecekteki çalışmalarda farklı stres şiddetleri ve uygulama sürelerinin, farklı aronya çeşitleri ve tarla koşulları altında değerlendirilmesiyle birlike kombine strese toleransın altında yatan moleküler ve metabolik mekanizmaların araştırılması önerilmektedir.

🔖

Cite This Article

Kaya, Ç., *, Ç. K. (2026). Effects of Combined Drought and Salinity Stress on Oxidative Damage and the Antioxidant Enzyme System in Aronia (Aronia melanocarpa L.) ‘Nero’ under In Vitro Conditions. Anadolu Tarım Bilimleri Dergisi, Cilt: 41. https://doi.org/10.7161/omuanajas.1900528

📄 Full Text PDF Viewer

📄

PDF görüntüleyici hazırlanıyor...

References (78)

  1. Aebi, H., 1984. Catalase in vitro. Methods in Enzymology, 105: 121-126. doi:10.1016/S0076-6879(84)05016-3.
  2. Aggarwal, J., Kaur, R., Mittal, A., Atri, C., Gupta, M., Sharma, A., 2025. Lipoxygenase 2 is a potential resistance determinant in Brassica juncea–B. fruticulosa derived lines against Lipaphis erysimi infestation. Acta Physiologiae Plantarum, 47(5): 54.
  3. Alptekin, B., Kunkowska, A.B., 2025. How plants adapt to combined and sequential abiotic stresses: A transcriptomics approach. Plant Physiology, 197(1): 1-2. doi:10.1093/plphys/kiaf006.
  4. Alvarez, M.E., Savouré, A., Szabados, L., 2022. Proline metabolism as regulatory hub. Trends in Plant Science, 27(1): 39-55.
  5. An, M., Huang, X., Long, Y., Wang, Y., Tan, Y., Qin, Z., Wang, Y., 2025. Salt tolerance evaluation and key salt-tolerant traits at germination stage of upland cotton. Frontiers in Plant Science, 15: 1489380.
  6. Angon, P.B., Tahjib-Ul-Arif, M., Samin, S.I., Habiba, U., Hossain, M.A., Brestic, M., 2022. How do plants respond to combined drought and salinity stress?—A systematic review. Plants, 11(21): 2884.
  7. Arora, D., Jain, P., Singh, N., Kaur, H., Bhatla, S.C., 2016. Mechanisms of nitric oxide crosstalk with reactive oxygen species scavenging enzymes during abiotic stress tolerance in plants. Free Radical Research, 50(3): 291-303.
  8. Badavath, H., Saski, C.A., 2026. Plant adaptation to salinity: Physiological pathways and prospects for crop improvement—A review. Genes, 17(3): 253.
  9. Barrs, H.D., Weatherley, P.E., 1962. A re-examination of the relative turgidity technique for estimating water deficits in leaves. Australian Journal of Biological Sciences, 15(3): 413-428. doi:10.1071/BI9620413.
  10. Bates, L.S., Waldren, R.P.A., Teare, I.D., 1973. Rapid determination of free proline for water-stress studies. Plant and Soil, 39(1): 205-207.
  11. Beauchamp, C., Fridovich, I., 1971. Superoxide dismutase: Improved assays and an assay applicable to acrylamide gels. Analytical Biochemistry, 44(1): 276-287.
  12. Belal, H.E., Abdelpary, M.A., Desoky, E.S.M., Ali, E.F., Al Kashgry, N.A.T., Rady, M.M., Sayed, A.A., 2023. Effect of eco-friendly application of bee honey solution on yield, physio-chemical, antioxidants, and enzyme gene expressions in excessive nitrogen-stressed common bean (Phaseolus vulgaris L.) plants. Plants, 12(19): 3435.
  13. Bradford, M.M., 1976. A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding. Analytical Biochemistry, 72(1-2): 248-254.
  14. Brand-Williams, W., Cuvelier, M.E., Berset, C., 1995. Use of a free radical method to evaluate antioxidant activity. LWT - Food Science and Technology, 28(1): 25-30. doi:10.1016/S0023-6438(95)80008-5.
  15. Brown, J.H., 2025. Applications of Archaeal Antioxidant Coenzyme M as an Agricultural Biostimulant. PhD Thesis. University of Nebraska-Lincoln, Department of Biochemistry, Lincoln, Nebraska.
  16. Chance, B., Maehly, A.C., 1955. Assay of catalases and peroxidases. Methods in Enzymology, 2: 764-775. doi:10.1016/S0076-6879(55)02300-8.
  17. Chauhan, P., Mir, R.A., Khah, M.A., 2022. Ascorbate–glutathione cycle: Nitric oxide and phytohormone interactions for plant stress tolerance. In: Ahanger, M.A., Ahmad, P. (Eds). Nitric Oxide in Plants. Wiley, pp. 148-178. doi:10.1002/9781119800156.ch8.
  18. Dixit, S., Sivalingam, P.N., Baskaran, R.M., Senthil-Kumar, M., Ghosh, P.K., 2024. Plant responses to concurrent abiotic and biotic stress: Unravelling physiological and morphological mechanisms. Plant Physiology Reports, 29(1): 6-17.
  19. Dubrovna, O.V., Mykhalska, S.I., Komisarenko, A.G., 2022. Using proline metabolism genes in plant genetic engineering. Cytology and Genetics, 56(4): 361-378.
  20. Ekinci, H., Şaşkın, N., Dikmetaş, B., Ak, B.E., 2024. Determination of the effects of drought stress on Aronia melanocarpa cv. Nero in vitro conditions. International Journal of Agriculture, Environment and Food Sciences, 8(4): 760-767.
  21. El Moukhtari, A., Cabassa, C., Durand, N., Farissi, M., Savouré, A., 2025. Exogenous proline supply improves growth, antioxidant defense system, and nutrient homeostasis in salt-stressed alfalfa (Medicago sativa L.). Journal of Soil Science and Plant Nutrition, 25(2): 3603-3614.
  22. El Moukhtari, A., Cabassa-Hourton, C., Crilat, E., Carol, P., Lamsaadi, N., Hidri, R., Savouré, A., 2023. Salt stress is alleviated by either proline or silicon but not by their combination in alfalfa (Medicago sativa L.) inoculated with a salt-tolerant Ensifer meliloti strain. Journal of Plant Growth Regulation, 42(7): 4048-4062.
  23. Ferreira, J.P., Vidal, M.S., Baldani, J.I., 2025. Exploring ACC deaminase-producing bacteria for drought stress mitigation in Brachiaria. Frontiers in Plant Science, 16: 1607697.
  24. Foyer, C.H., Halliwell, B., 1976. The presence of glutathione and glutathione reductase in chloroplasts: A proposed role in ascorbic acid metabolism. Planta, 133(1): 21-25.
  25. Foyer, C.H., Kunert, K., 2024. The ascorbate-glutathione cycle coming of age. Journal of Experimental Botany, 75(9): 2682-2699.
  26. George, E.F., Hall, M.A., De Klerk, G.J., 2008. Plant Propagation by Tissue Culture. Volume 1: The Background. 3rd ed. Springer, Dordrecht, 502 p.
  27. Gupta, A., Senthil-Kumar, M., 2017. Concurrent stresses are perceived as new state of stress by the plants: Overview of impact of abiotic and biotic stress combinations. In: Senthil-Kumar, M. (Ed). Plant Tolerance to Individual and Concurrent Stresses. Springer, New Delhi, pp. 1-15. doi:10.1007/978-81-322-3706-8_1.
  28. Hajam, Y.A., Lone, R., Kumar, R., 2023. Role of plant phenolics against reactive oxygen species (ROS) induced oxidative stress and biochemical alterations. In: Plant Phenolics in Abiotic Stress Management. Springer Nature Singapore, pp. 125-147. doi:10.1007/978-981-19-6426-8_7.
  29. Hasanuzzaman, M., Fujita, M., 2022. Plant oxidative stress: Biology, physiology and mitigation. Plants, 11(9): 1185. Heath, R.L., Packer, L., 1968. Photoperoxidation in isolated chloroplasts: I. Kinetics and stoichiometry of fatty acid peroxidation. Archives of Biochemistry and Biophysics, 125(1): 189-198. doi:10.1016/0003-9861(68)90654-1.
  30. Jiang, Z., van Zanten, M., Sasidharan, R., 2025. Mechanisms of plant acclimation to multiple abiotic stresses. Communications Biology, 8(1): 655.
  31. Jurendić, T., Ščetar, M., 2021. Aronia melanocarpa products and by-products for health and nutrition: A review. Antioxidants, 10(7): 1052.
  32. Kissoudis, C., Sunarti, S., Van De Wiel, C., Visser, R.G., van der Linden, C.G., Bai, Y., 2016. Responses to combined abiotic and biotic stress in tomato are governed by stress intensity and resistance mechanism. Journal of Experimental Botany, 67(17): 5119-5132.
  33. Kulling, S.E., Rawel, H.M., 2008. Chokeberry (Aronia melanocarpa)—A review on the characteristic components and potential health effects. Planta Medica, 74(13): 1625-1634.
  34. Lawlor, D.W., 1970. Absorption of polyethylene glycols by plants and their effects on plant growth. New Phytologist, 69(2): 501-513.
  35. Mahmood, M.S.M., Pırlak, L., 2025. Salt stress sensitivity of chokeberry (Aronia melanocarpa L.) in vitro and in vivo conditions. Selcuk Journal of Agriculture and Food Sciences, 39(1): 31-41. doi:10.15316/SJAFS.2025.004.
  36. Maiti, I., Dadarwal, B.K., Saini, D.R., Gupta, S.K., Yadav, N., Prakash, P., 2025. Impact of elevated CO2 and temperature on plant redox metabolism. In: Climate Change: Impact of Elevated CO2 and Temperature on Crops, Weeds and Soil Microbes. CRC Press, pp. 159-173. doi:10.1201/9781003483854-14.
  37. Martins, J.P.R., Santos, E.R., Rodrigues, L.C.A., Gontijo, A.B.P.L., Falqueto, A.R., 2018. Effects of 6-benzylaminopurine on photosystem II functionality and leaf anatomy of in vitro cultivated Aechmea blanchetiana. Biologia Plantarum, 62(4): 793-800.
  38. Melino, V., Tester, M., 2023. Salt-tolerant crops: Time to deliver. Annual Review of Plant Biology, 74(1): 671-696. Michel, B.E., Kaufmann, M.R., 1973. The osmotic potential of polyethylene glycol 6000. Plant Physiology, 51(5): 914-916.
  39. Mishra, G., 2026. Biochemical memory in plants: A missing component in climate resilience models. Journal of Plant Physiology, 318: 154728. doi:10.1016/j.jplph.2026.154728.
  40. Mishra, N., Jiang, C., Chen, L., Paul, A., Chatterjee, A., Shen, G., 2023. Achieving abiotic stress tolerance in plants through antioxidative defense mechanisms. Frontiers in Plant Science, 14: 1110622.
  41. Mittler, R., Karlova, R., Bassham, D.C., Lawson, T., 2025. Crops under stress: Can we mitigate the impacts of climate change on agriculture and launch the ‘Resilience Revolution’? Philosophical Transactions of the Royal Society B: Biological Sciences, 380: 1927.
  42. Mittler, R., Zandalinas, S.I., Fichman, Y., Van Breusegem, F., 2022. Reactive oxygen species signalling in plant stress responses. Nature Reviews Molecular Cell Biology, 23(10): 663-679.
  43. Molodchenkova, O., Fanin, Y., 2025. Phenolic metabolism, antioxidant processes, and lipid metabolism in wheat and barley plants: Role in protective responses against fungal phytopathogens. Publishing House “Baltija Publishing”.
  44. Murashige, T., Skoog, F., 1962. A revised medium for rapid growth and bio assays with tobacco tissue cultures. Physiologia Plantarum, 15(3): 473-497. doi:10.1111/j.1399-3054.1962.tb08052.x.
  45. Nagar, A., Guru, A., Tripathi, A., 2026. Abiotic stress management in Brassica napus. In: Crop Improvement Strategies in Brassica Species: Stress Resistance Strategies, Volume 2. Springer Nature, pp. 1-19. doi:10.1007/978-981-95-4713-5_1.
  46. Nakano, Y., Asada, K., 1981. Hydrogen peroxide is scavenged by ascorbate-specific peroxidase in spinach chloroplasts. Plant and Cell Physiology, 22(5): 867-880.
  47. Nas, Z., Eşitken, A., Pırlak, L., 2023. The responses of ‘Viking’ aronia variety to salinity stress under in vitro conditions. Erwerbs-Obstbau, 65(6): 2547-2552.
  48. Noctor, G., Cohen, M., Trémulot, L., Châtel-Innocenti, G., Van Breusegem, F., Mhamdi, A., 2024. Glutathione: A key modulator of plant defence and metabolism through multiple mechanisms. Journal of Experimental Botany, 75(15): 4549-4572.
  49. Nowicka, B., Hoque, T.S., Khatun, S.M., Naim, J., Hasan, A.K., Hossain, M.A., 2023. Heavy-metal-induced reactive oxygen species and methylglyoxal formation and detoxification in crop plants: Modulation of tolerance by exogenous chemical compounds. In: Heavy Metal Toxicity and Tolerance in Plants: A Biological, Omics, and Genetic Engineering Approach. pp. 461-492. doi:10.1002/9781119906506.ch21.
  50. Okumuş, O., Şimşek, Ö., Isak, M.A., Şahin, N.K., Aydin, A., Eren, B., Yaman, M., 2025. Integrating machine learning and in vitro screening to evaluate drought and temperature stress responses for Vicia species. Processes, 13(6): 1845.
  51. Oliwa, J., Sieprawska, A., Dyba, B., 2026. Nighttime warming enhances photosynthetic activity and induces changes in chloroplast membrane structure and antioxidant profile in Platycerium ferns. Scientific Reports, 16: 5976. doi:10.1038/s41598-026-37176-9.
  52. Ou, T., Jiang, K., Wang, L., Liu, X., Zhang, K., Wen, J., Xie, J., 2026. Rhizobacterial coordination of reactive oxygen species homeostasis underpins mulberry resilience to waterlogging. Plant Physiology and Biochemistry, 111085.
  53. Parvin, K., Nahar, K., Mohsin, S.M., Al Mahmud, J., Fujita, M., Hasanuzzaman, M., 2022. Plant phenolic compounds for abiotic stress tolerance. In: Managing Plant Production Under Changing Environment. Springer Nature Singapore, pp. 193-237. doi:10.1007/978-981-16-5059-8_8.
  54. Parween, G., Akram, S., Singh, T., Saema, S., Khatoon, K., Rahman, L.U., 2025. Advances in elicitor-mediated secondary metabolite production in plant cell suspensions: Factors and mechanisms. In: Elicitation Methods for Improvement in Plant Secondary Metabolites. Springer Nature, pp. 57-83.
  55. Rane, J., Singh, A.K., Tiwari, M., Prasad, P.V., Jagadish, S.K., 2022. Effective use of water in crop plants in dryland agriculture: Implications of reactive oxygen species and antioxidative system. Frontiers in Plant Science, 12: 778270.
  56. Ray, S.R., Roy, A., 2025. Plants under salinity stress: A physiological and antioxidant resistance scheme. Journal of Biochemistry, 11(1). doi:10.52338/job.2025.4728.
  57. Ray, S.R., Majid, A., Waza, S.A., Islam, A.K.M.A., 2025. Plant breeding approaches for enhancing abiotic stress tolerance in crops: A review. Madras Agricultural Journal, 112(3): 11-22. doi:10.29321/MAJ.10.90JE29.
  58. Sachdev, S., Ansari, S.A., Ansari, M.I., Fujita, M., Hasanuzzaman, M., 2021. Abiotic stress and reactive oxygen species: Generation, signaling, and defense mechanisms. Antioxidants, 10(2): 277.
  59. Sánchez-Bermúdez, M., Del Pozo, J.C., Pernas, M., 2022. Effects of combined abiotic stresses related to climate change on root growth in crops. Frontiers in Plant Science, 13: 918537.
  60. Sathi, K.S., Masud, A.A.C., Anee, T.I., Rahman, K., Ahmed, N., Hasanuzzaman, M., 2022. Soybean plants under waterlogging stress: Responses and adaptation mechanisms. In: Managing Plant Production Under Changing Environment. Springer Nature Singapore, pp. 103-134. doi:10.1007/978-981-16-5059-8_5.
  61. Shajar, F., Nabi, A.U., Manzoor, A., Saleem, S., Mushtaq, N.U., Rehman, R.U., 2025. Oxidative stress generation and antioxidant defense response in plants during abiotic stress. In: Plant Biotechnology and Sustainable Agriculture: Bridging the Gap for Global Food Security. pp. 205-220.
  62. Sharaya, R., Tuteja, N., Gill, R., Gill, S.S., 2023. Reactive oxygen species (ROS) scavenging and methylglyoxal (MG) detoxification confers salinity tolerance in mustard cultivars inoculated with Piriformospora (Serendipita) indica. South African Journal of Botany, 163: 630-645.
  63. Shehzad, J., Mustafa, G., 2023. Mechanism of reactive oxygen species regulation in plants. In: Faizan, M., Hayat, S., Ahmed, S.M. (Eds). Reactive Oxygen Species: Prospects in Plant Metabolism. Springer Singapore, pp. 17-41.
  64. Sidor, A., Gramza-Michałowska, A., 2019. Black chokeberry Aronia melanocarpa L.—A qualitative composition, phenolic profile and antioxidant potential. Molecules, 24(20): 3710.
  65. Singleton, V.L., Rossi, J.A., 1965. Colorimetry of total phenolics with phosphomolybdic-phosphotungstic acid reagents. American Journal of Enology and Viticulture, 16(3): 144-158. doi:10.5344/ajev.1965.16.3.144.
  66. Sinha, R., Peláez-Vico, M.Á., Dhakal, S., Ghani, A., Myers Jr, R., Verma, M., Mittler, R., 2025. Transcriptomic and physiological responses of soybean plants subjected to a combination of water deficit and heat stress under field conditions. bioRxiv, 2025-05.
  67. Sood, M., 2025. Reactive oxygen species (ROS): Plant perspectives on oxidative signalling and biotic stress response. Discover Plants, 2(1): 187.
  68. Swain, B., Sahoo, L., Tiwari, V., Kumar, C., Joshi, N.C., Singh, P.K., Yadav, D., 2025. Reactive oxygen species. In: Soil Health and Nutrition Management, p. 169.
  69. Thakur, K., Garg, N., 2022. Oxidative stress and antioxidant enzymes in cereals under abiotic stress. In: Sustainable Remedies for Abiotic Stress in Cereals. Springer Nature Singapore, pp. 51-82. doi:10.1007/978-981-19-5121-3_3.
  70. Toyokuni, S., Kong, Y., Mi, D., 2022. On ‘Photoperoxidation in isolated chloroplasts: I. Kinetics and stoichiometry of fatty acid peroxidation’ by Robert L. Heath and Lester Packer. Archives of Biochemistry and Biophysics, 726: 109133.
  71. ul Islam, S.N., Asgher, M., Khan, N.A., 2023. Hydrogen peroxide and its role in abiotic stress tolerance in plants. In: Gasotransmitters Signaling in Plant Abiotic Stress. Springer International Publishing, Cham, pp. 167-195. doi:10.1007/978-3-031-30858-1_9.
  72. Velikova, V., Yordanov, I., Edreva, A., 2000. Oxidative stress and some antioxidant systems in acid rain-treated bean plants: Protective role of exogenous polyamines. Plant Science, 151(1): 59-66. doi:10.1016/S0168-9452(99)00197-1.
  73. Wagay, N.A., Rafiq, S., Khan, A., Kaloo, Z.A., Malik, A.R., Pulate, P.V., 2023. Impact of phenolics on drought stress and expression of phenylpropanoid pathway genes. In: Plant Phenolics in Abiotic Stress Management. Springer Nature Singapore, pp. 265-285. doi:10.1007/978-981-19-6426-8_13.
  74. Wu, Y., Bai, L., Dai, X., Ba, L., Wan, J., Liang, W., Fan, Z., 2025. Comparative transcriptomic analysis reveals the reactive oxygen species metabolism involving in melatonin-alleviated chilling injury in postharvest banana fruit. Plant Physiology and Biochemistry, 222: 109693.
  75. Yadav, P., Ansari, M.W., Saini, S., Punia, S., Kaula, B.C., Rani, V., Tuteja, N., 2024. Review and future prospects on the impact of abiotic stresses and tolerance strategies in medicinal and aromatic plants. Brazilian Journal of Botany, 47(3): 683-701.
  76. Zandalinas, S.I., Balfagón, D., Gómez-Cadenas, A., Mittler, R., 2022. Responses of plants to climate change: Metabolic changes during abiotic stress combination in plants. Journal of Experimental Botany, 73(11): 3339-3354.
  77. Zandi, P., Schnug, E., 2022. Reactive oxygen species, antioxidant responses and implications from a microbial modulation perspective. Biology, 11(2): 155.
  78. Zhu, D., Zhang, X., Wang, X., Li, X., Prusky, D., Bi, Y., Zong, Y., 2026. Deletion of PeCdc42 reduces Penicillium expansum pathogenicity in apple fruit by modulating host reactive oxygen species metabolism. Postharvest Biology and Technology, 234: 114076.