Forest Research and Development

Forest Research and Development

Antioxidant enzyme responses of two-year-old Pistacia atlantica Desf. seedlings to combined drought and dust Stress

Document Type : Scientific article

Authors
Fatemeh Pourfallahi 1
Babak Pilehvar 2
Zahra Mirazadi 3
Seyed Vahid Sayedena 3
1 PhD Student of Forestry, Faculty of Natural Resources, Lorestan University, Lorestan, I. R. Iran
2 Professor, Forestry Department, Faculty of Natural Resources, Lorestan University, Lorestan, I. R. Iran
3 Assistant Professor, Forestry Department, Faculty of Natural Resources, Lorestan University, Lorestan, I. R. Iran
10.30466/jfrd.2026.56637.1785
Abstract
Background and Objective: Over the past two decades, the severity of drought and the frequency of dust storms have increased markedly in Iran, particularly in the western and southwestern provinces, imposing substantial pressure on natural ecosystems. The Zagros forests, one of the country's most valuable and vulnerable forest ecosystems, are highly susceptible to the combined impacts of drought and dust deposition. These environmental stressors reduce tree growth, disrupt physiological processes, and accelerate forest decline. Wild pistachio (Pistacia atlantica Desf.), an ecologically and economically important tree species of Iran's arid and semi-arid regions, plays a crucial role in maintaining ecosystem stability. Therefore, this study aimed to investigate the combined effects of drought and dust stress on the antioxidant enzyme activities of wild pistachio seedlings in order to better understand their defense mechanisms and provide insights for improving their resilience to environmental stress.
Material and Methods: This study was conducted from March 2023 to December 2023 in the greenhouse of the Faculty of Agriculture and Natural Resources, Lorestan University. A factorial experiment was arranged as a split-plot design within a randomized complete block design using 384 wild pistachio seedlings. Drought stress was applied at three irrigation levels corresponding to 100% (control), 75%, and 50% of field capacity, while dust stress consisted of four concentrations: 0, 350, 750, and 1,500 μg m⁻³. Growth chambers with transparent plastic covers were constructed inside the greenhouse, and the seedlings were placed within these chambers. The potting medium consisted of equal proportions of agricultural soil, sand, and farmyard manure, similar to the substrate commonly used in forest nurseries. At the end of the experiment, leaf samples were collected and the activities of catalase (CAT), peroxidase (POD), superoxide dismutase (SOD), and ascorbate peroxidase (APX) were determined under laboratory conditions.
Results: Drought and dust stresses significantly affected the activities of all antioxidant enzymes at the 1% probability level. However, the interaction between the two stressors was not significant for APX activity, indicating that each stress factor independently influenced this enzyme. Catalase activity increased markedly with increasing stress intensity. Under severe drought (50% of field capacity), APX and SOD activities showed a slight decline at the initial dust concentration (350 μg m⁻³), which may reflect temporary impairment of the antioxidant defense system under oxidative stress. With further increases in dust concentration to 750 and 1,500 μg m⁻³, SOD activity increased substantially, suggesting activation of compensatory defense mechanisms to counteract reactive oxygen species. Although POD activity was significantly influenced by both drought and dust stress, its response did not exhibit a consistent increasing or decreasing trend.
Conclusion: Wild pistachio seedlings responded to the combined effects of drought and dust stress by significantly enhancing the activities of CAT, POD, and SOD, demonstrating an effective antioxidant defense mechanism. The marked increase in SOD activity, as the first line of defense against reactive oxygen species, highlights its critical role in protecting cellular integrity and enhancing stress tolerance. Nevertheless, under severe drought and dust conditions, physiological performance and seedling growth were substantially impaired, increasing the likelihood of seedling mortality. Consequently, mitigating dust sources through wetland restoration, securing environmental water allocations, establishing stress-tolerant vegetation in dust-emitting areas, and strengthening regional cooperation for transboundary dust control and sustainable water resource management are recommended to reduce environmental stress and improve the resilience of Zagros forest ecosystems.
Keywords
Peroxidase
catalase
superoxide dismutase
Zagros forests
Subjects
Aebi, H. Catalase in vitro. Methods in Enzymology 1984, 105, 121–126.
Ahmadi, M.; Ranjbar, G.; Etemadi, N. Antioxidant responses of Quercus seedlings under combined drought and dust stress. Journal of Forestry Research 2020, 31(4), 1321–1330.
Ahmadi, S.; Khan, M. A.; Iqbal, M. Response of Quercus species to drought stress: Antioxidant enzymes and physiological changes. Journal of Plant Physiology 2019, 234, 12–20. https://doi.org/10.1016/j.jplph.2019.02.004.
Amini, Z.; Moalemi, N.; Saadati, S. Effect of Water Deficit on Proline Content and Activity of Antioxidant Enzymes among Three Olive (Olea europaea L.) Cultivars. Journal of Plant Research 2014, 27 (2), 156–167. (In Persian)
Chance, B.; Maehly, A. C. Assay of catalases and peroxidases. Methods in Enzymology 1955, 2, 764–775.
Esmaili, A.; Mousavi Mirkala, S. R.; Alijanpour, A.; Hajarian, M.; Ghanbari, S. The effect of altitude on quantitative and qualitative characteristics and estimation of Pistacia atlantica fruits in the forest stands in Sardasht, West Azerbaijan Province. Journal of Forest Research and Development 2022, 7(4), 653–670. (In Persian).
Fernández-Marín, B.; Casas, R.; Ancín, A. Antioxidant enzyme activity in Fraxinus excelsior under drought conditions. Plant Biology 2013, 15(5), 819–826. https://doi.org/10.1111/plb.12032.
Gill, S. S.; Tuteja, N. Reactive oxygen species and antioxidant machinery in abiotic stress tolerance in crop plants. Plant Physiology and Biochemistry 2010, 48(12), 909–930. https://doi.org/10.1016/j.plaphy.2010.08.016
Harb, A.; Samarah, N. Gene expression and activity of antioxidant enzymes in barley (Hordeum vulgare L.) under controlled severe drought. Journal of Agronomy and Crop Science 2015.
Hashempour, F.; Rostami Shahraji, T.; Assareh, M. H.; Shariat, A. Impact of Drought Stress on Some Physiological Traits in Five Eucalypt Species. Iranian Journal of Forest and Poplar Research 2011, 19 (2), 222–233. (In Persian)
 
IPCC. Climate Change 2021: The Physical Science Basis; Cambridge University Press: Cambridge, 2021.
Iranmanesh, Y.; Jahanbazy, H.; Talebi, M.; Mahinpour, H. Effect of morphological variables, altitude and tree gender on gum production of Pistacia atlantica in Chaharmahal & Bakhtiari Province forests. Journal of Forest Research and Development 2019, 5(2), 195–207. (In Persian).
Jafaryan, E.; Pilehvar, B.; Tavakli, M. Physiological responses of mature Persian oak (Quercus brantii L.) under natural conditions to drought stress. Forest Research and Development 2024, 10(2), 167–181. (In Persian).
Karimi, N.; Hassani, A.; Mohammadi, H. Drought tolerance mechanisms in Pistacia atlantica seedlings. Journal of Arid Environments 2019, 164, 58–66.
Kazemi, H.; Ahmadi, F.; Nasrollahi, S. Effect of dust stress on antioxidant enzyme activities in tree seedlings under drought conditions. Iranian Journal of Plant Physiology 2017, 7(4), 1913–1922.
Khan, M. I. R.; et al. Plant responses to dust pollution: Impact on photosynthesis and antioxidant metabolism. Environmental Science and Pollution Research 2017, 24, 21128–21140. https://doi.org/10.1007/s11356-017-9749-7.
Kozlowski, T. T. Responses of woody plants to flooding and salinity. Tree Physiology 2011, 31(6), 567–574.
Li, X.; Zhang, J.; Huang, Y. Response of antioxidant enzymes to drought stress in Populus deltoides seedlings. Plant Science 2012, 197, 79–86. https://doi.org/10.1016/j.plantsci.2012.09.004.
Mahajan, S.; Tuteja, N. Cold, salinity and drought stresses: An overview. Archives of Biochemistry and Biophysics 2005, 444(2), 139–158.
Middleton, N. Desert dust hazards: A global review. Aeolian Research 2017, 24, 53–63. https://doi.org/10.1016/j.aeolia.2016.12.001.
Mirzaei, J.; Karamshahi, A. A. The effect of drought stress on the growth and some physiological characteristics of Pistacia atlantica Desf. seedlings. Wood and Forest Science and Technology Research Journal 2014, 22(1). (In Persian).
Misra, H. P.; Fridovich, I. The generation of superoxide radical during auto oxidation. Journal of Biological Chemistry 1972, 247, 6960–6966.
Mittler, R. Oxidative stress, antioxidants and stress tolerance. Trends in Plant Science 2002, 7(9), 405–410.
Mittler, R.; Vanderauwera, S.; Gollery, M.; Van Breusegem, F. Reactive Oxygen Gene Network of Plants. Trends in Plant Science 2004, 9 (10), 490–498.
Mokhtar, M.; El-Kholy, A.; El-Bebany, A. Effect of drought stress on antioxidant enzymes in Acacia nilotica seedlings. Environmental and Experimental Botany 2014, 102, 1–8. https://doi.org/10.1016/j.envexpbot.2014.03.005.
Moradi, F.; Ghorbani, R. Antioxidant enzyme activities in Pistacia atlantica under drought and pollution stress. Iranian Journal of Forestry and Range Management 2018, 72(1), 45–56. (In Persian).
Morales, D.; García-Criado, B.; Retana, J. Antioxidant response in Quercus ilex seedlings under drought stress. Environmental and Experimental Botany 2015, 111, 119–127. https://doi.org/10.1016/j.envexpbot.2014.09.009.
Naji, H. R.; Roushani Niaa, F.; Tongo, A.; Soheili, F.; Arminian, A. Effect of simulated dust storm conditions on the physiological features of wild pistachio. Forests 2024, 20(1), 16–24. https://doi.org/10.1080/21580103.2023.2280647.
Nakano, Y.; Asada, K. Hydrogen peroxide is scavenged by ascorbate-specific peroxidase in spinach chloroplasts. Plant and Cell Physiology 1981, 22(5), 867–880.
Nawaz, M. F.; Rashid, M. H. U.; Saeed-Ur-Rehman, M.; Gul, S.; Farooq, T. H.; Sabir, M. A.; Iftikhar, J.; Abdelsalam, N. R.; Dessoky, E. S.; Alotaibi, S. S. Effect of dust types on the eco-physiological response of three tree species seedlings: Eucalyptus camaldulensis, Conocarpus erectus and Bombax ceiba. Atmosphere 2022, 13(7), 1010, 1–12. https://doi.org/10.3390/atmos13071010.
Pandey, P.; et al. Impact of combined abiotic stresses on oxidative damage and antioxidant defense in plants. 2015.
Pérez-López, U.; Gómez-Cadenas, A.; Muñoz-Rueda, A. Drought-induced changes in antioxidant enzymes in Pinus halepensis seedlings. Tree Physiology 2009, 29(9), 1173–1181. https://doi.org/10.1093/treephys/tpp059.
Saeidi Abueshaghi, Z.; Pilehvar, B.; Sayedena, S. V. Effect of drought stress on morphophysiological and biochemical traits of Cercis siliquastrum L. seedlings. Iranian Journal of Forest and Poplar Research 2021, 29(1), 91–100. https://doi.org/10.22059/jfrd.2021.121376. (In Persian).
Shahrokhi, S.; Tavakoli, E.; Moradi, F. Oxidative stress and antioxidant enzyme activity in plants exposed to particulate matter pollution. Ecotoxicology and Environmental Safety 2018, 157, 228–235.
Shahsavani, A.; et al. The evaluation of PM10, PM2.5, and PM1 concentrations during Middle Eastern dust storms in Ahvaz, Iran. Environmental Monitoring and Assessment 2020, 192, 206. https://doi.org/10.1007/s10661-020-8155-9.
Sharma, P.; Dubey, R. S.; Chauhan, P. S. Impact of combined drought and dust stress on antioxidant enzymes in Eucalyptus seedlings. Environmental Science and Pollution Research 2019, 26(21), 21800–21812.
Singh, R.; Kumar, P. Role of peroxidase and catalase enzymes in oxidative stress tolerance of plants. Plant Biology Journal 2017, 19(3), 265–273. https://doi.org/10.1111/plb.12543.
Suzuki, N.; Rivero, R. M.; Shulaev, V.; Blumwald, E.; Mittler, R. Abiotic and biotic stress combinations. New Phytologist 2014, 203(1), 32–43. https://doi.org/10.1111/nph.12797.
Wali, A.; Khamishi, S.; Mousavi, S. H.; Panahi, F.; Temski, A. Climatic analysis and tracking of dust plumes in the south and center of Iran. Journal of Environmental Science 2014, 40, 961–972. (In Persian).
Xiong, S.; Wang, Y.; Chen, Y.; Gao, M.; Zhao, Y.; Wu, L. Effects of drought stress and rehydration on physiological and biochemical characteristics of four oak species in China. Plants 2022, 11, 679. https://doi.org/10.3390/plants11050679.
Yaghmaei, L.; Jafari, R.; Soltani, S.; Eshghizadeh, H. R.; Jahanbazy, H. Interaction effects of dust and water deficit stresses on growth and physiology of Persian oak (Quercus brantii Lindl.). Journal of Sustainable Forestry 2022, 41(2), 134–158.
Yousefvand, P.; Pilehvar, B.; Nasrolahi, A. H. Morphological, physiological, and biochemical responses of Pistacia atlantica seedlings to elevated CO₂ concentration and drought stress. European Journal of Forest Research 2023. https://doi.org/10.1007/s10342-023-01548-x.
Zhang, L.; Wang, H.; Liu, Y. Combined effects of drought and air pollution on antioxidant enzymes in tree seedlings. Environmental Pollution 2018, 242, 874–882.
Zoljoodi, M.; Didevarasl, A. Dust events in the western parts of Iran and the relationship with drought expansion over the dust-source areas in Iraq and Syria. Atmospheric and Climate Sciences 2013, 3, 321–336. https://doi.org/10.4236/acs.2013.33034
Forest Research and Development
Volume 12, Issue 2
Summer 2026
Pages 203-218