Research in Plant Metabolites

Research in Plant Metabolites

Evaluation of Seed Priming on Quinoa (Chenopodium quinoa Willd.) Germination Behavior and Enzymatic Activities Under Lead Nitrate Stress Conditions

Document Type : Original Article

Authors
Fatemeh Jahanbakhshi 1
Alireza Abdali Mashhadi
Seyed Amir Moosavi 2
Amin Lotfi Jalal Abadi 1
1 Department of Production and Plant Genetics Engineering, Agricultural Sciences and Natural Resources University of Khuzestan, Mollasani, Iran
2 Department of Production and Plant Genetics Engineering, Agricultural Sciences and Natural Resources University of Khuzestan, Mollasani, Iran.
10.22034/jrpsm.2026.583542.1115
Abstract
Background and Objective: In recent decades, the increasing concentration of heavy metals in agricultural soils due to industrial and anthropogenic activities has emerged as a major environmental challenge. Lead (Pb), as one of the most persistent and toxic heavy metals, exerts destructive effects on early growth stages, particularly germination and seedling establishment, by inducing oxidative stress, disrupting water uptake, and impairing metabolic activities. Quinoa, a pseudo-cereal with high nutritional value and resilience to adverse conditions, holds great promise for cultivation in stressed environments; however, it remains sensitive to Pb toxicity during its early growth phases. Seed priming technology represents a modern and cost-effective strategy to address this challenge. Despite various studies, the simultaneous efficiency of different iron sources (especially nanoparticles) and calcium in mitigating Pb toxicity in Quinoa has received limited attention. Therefore, this study aimed to evaluate and compare the effects of priming with nano-iron, iron chloride, and calcium sulfate on germination indices, antioxidant defense system activity, and Pb accumulation in Quinoa seedlings.
Materials and Methods: This factorial experiment, based on a completely randomized design with three replications, was conducted in 2024 at the Seed Technology Laboratory of Khuzestan University. Priming treatments included iron chloride (FeCl₃) and nano-iron at 0, 4, 6, 8, and 10 mg/kg, and calcium sulfate (CaSO₄) at 0, 0.5, 1, 1.5, and 2 mM. Seeds were immersed for 6 hours. After priming, seeds were exposed to Pb nitrate stress levels (0, 20, 40, 80, and 100 mg/kg) in a germinator. Germination was monitored daily for one week. Morphological parameters (radicle and plumule length) were measured using Digimizer software. Standard indices, germination percentage (GP), germination rate (GR), mean germination time (MGT), and seedling vigor index (SVI) were calculated. Antioxidant enzyme activities (catalase, CAT; peroxidase, POD) were measured spectrophotometrically. Pb accumulation in tissues was determined via atomic absorption spectrophotometry after acid digestion. Statistical analysis used SAS software, with means compared using LSD test (P<0.05). The slicing method analyzed interaction effects at different stress levels.
Findings: Analysis of variance results indicated that all studied indices were significantly affected by both Pb stress and priming treatments. Increased lead nitrate concentrations led to a sharp decline in GP, GR, and SVI, while significantly increasing MGT. Conversely, priming treatments neutralized a major portion of these inhibitory effects. at 1–2 mM was the superior treatment for improving germination indices, maintaining a germination percentage of 93% even under the highest Pb stress level. Regarding morphological traits, iron priming exhibited higher efficiency. Nano-iron at 6 mg/kg had the most significant impact on enhancing plumule and radicle length under mild and moderate stress, while at 10 mg/kg performed better in maintaining seedling vigor under severe stress conditions. Biochemical assessments revealed that as Pb stress intensified, the activities of CAT and POD increased as a defense mechanism. Priming treatments (especially 10 mg/kg and 2 mM ) significantly boosted these enzyme activities compared to the non-primed control, contributing to reduced oxidative damage and enhanced cell membrane stability. Another crucial finding was the impact of priming on heavy metal content; atomic absorption analysis demonstrated that priming with 10 mg/kg reduced Pb accumulation in seedling tissues by approximately 48% at the highest stress level. This phenomenon maybe results from the competitive inhibition between iron and lead ions at uptake sites and improved membrane integrity in primed seeds.
Conclusion: This research confirmed that while lead stress imposes severe constraints on the early establishment of quinoa, seed priming with calcium and iron elements serves as an efficient strategy to mitigate this toxicity. Calcium sulfate not only improves primary physical and enzymatic processes through calcium ions but also contributes to lead detoxification via its sulfate moiety; by providing the essential sulfur foundation required to strengthen the antioxidant defense system, it effectively ensures optimal germination speed and percentage. Conversely, iron priming (particularly nano-iron at low doses and iron chloride at high doses) stimulates seedling growth and enhances seed vigor. Notably, exhibited a distinct superiority in promoting enzymatic activities (catalase and peroxidase), demonstrating significant efficiency in boosting the antioxidant defense system and restricting the excessive entry of lead into plant tissues. Consequently, the application of these priming treatments is recommended for improving quinoa establishment in soils contaminated with heavy metals
Background and Objective: In recent decades, the increasing concentration of heavy metals in agricultural soils due to industrial and anthropogenic activities has emerged as a major environmental challenge. Lead (Pb), as one of the most persistent and toxic heavy metals, exerts destructive effects on early growth stages, particularly germination and seedling establishment, by inducing oxidative stress, disrupting water uptake, and impairing metabolic activities. Quinoa, a pseudo-cereal with high nutritional value and resilience to adverse conditions, holds great promise for cultivation in stressed environments; however, it remains sensitive to Pb toxicity during its early growth phases. Seed priming technology represents a modern and cost-effective strategy to address this challenge. Despite various studies, the simultaneous efficiency of different iron sources (especially nanoparticles) and calcium in mitigating Pb toxicity in Quinoa has received limited attention. Therefore, this study aimed to evaluate and compare the effects of priming with nano-iron, iron chloride, and calcium sulfate on germination indices, antioxidant defense system activity, and Pb accumulation in Quinoa seedlings.
Materials and Methods: This factorial experiment, based on a completely randomized design with three replications, was conducted in 2024 at the Seed Technology Laboratory of Khuzestan University. Priming treatments included iron chloride (FeCl₃) and nano-iron at 0, 4, 6, 8, and 10 mg/kg, and calcium sulfate (CaSO₄) at 0, 0.5, 1, 1.5, and 2 mM. Seeds were immersed for 6 hours. After priming, seeds were exposed to Pb nitrate stress levels (0, 20, 40, 80, and 100 mg/kg) in a germinator. Germination was monitored daily for one week. Morphological parameters (radicle and plumule length) were measured using Digimizer software. Standard indices, germination percentage (GP), germination rate (GR), mean germination time (MGT), and seedling vigor index (SVI) were calculated. Antioxidant enzyme activities (catalase, CAT; peroxidase, POD) were measured spectrophotometrically. Pb accumulation in tissues was determined via atomic absorption spectrophotometry after acid digestion. Statistical analysis used SAS software, with means compared using LSD test (P<0.05). The slicing method analyzed interaction effects at different stress levels.
Findings: Analysis of variance results indicated that all studied indices were significantly affected by both Pb stress and priming treatments. Increased lead nitrate concentrations led to a sharp decline in GP, GR, and SVI, while significantly increasing MGT. Conversely, priming treatments neutralized a major portion of these inhibitory effects. at 1–2 mM was the superior treatment for improving germination indices, maintaining a germination percentage of 93% even under the highest Pb stress level. Regarding morphological traits, iron priming exhibited higher efficiency. Nano-iron at 6 mg/kg had the most significant impact on enhancing plumule and radicle length under mild and moderate stress, while at 10 mg/kg performed better in maintaining seedling vigor under severe stress conditions. Biochemical assessments revealed that as Pb stress intensified, the activities of CAT and POD increased as a defense mechanism. Priming treatments (especially 10 mg/kg and 2 mM ) significantly boosted these enzyme activities compared to the non-primed control, contributing to reduced oxidative damage and enhanced cell membrane stability. Another crucial finding was the impact of priming on heavy metal content; atomic absorption analysis demonstrated that priming with 10 mg/kg reduced Pb accumulation in seedling tissues by approximately 48% at the highest stress level. This phenomenon maybe results from the competitive inhibition between iron and lead ions at uptake sites and improved membrane integrity in primed seeds.
Conclusion: This research confirmed that while lead stress imposes severe constraints on the early establishment of quinoa, seed priming with calcium and iron elements serves as an efficient strategy to mitigate this toxicity. Calcium sulfate not only improves primary physical and enzymatic processes through calcium ions but also contributes to lead detoxification via its sulfate moiety; by providing the essential sulfur foundation required to strengthen the antioxidant defense system, it effectively ensures optimal germination speed and percentage. Conversely, iron priming (particularly nano-iron at low doses and iron chloride at high doses) stimulates seedling growth and enhances seed vigor. Notably, exhibited a distinct superiority in promoting enzymatic activities (catalase and peroxidase), demonstrating significant efficiency in boosting the antioxidant defense system and restricting the excessive entry of lead into plant tissues. Consequently, the application of these priming treatments is recommended for improving quinoa establishment in soils contaminated with heavy metals
Keywords
Seedling establishment
Oxidative stress
Atomic absorption
Heavy metals
Catalaz
Ion toxicity
Abdoli, M. (2020). The effect of seed aging and hydro-priming on germination indices and the activity of some antioxidant enzymes in maize hybrid (Zea mays L.). Iranian Journal of Seed Science and Research, 7(2), 147-159. [In Persian].
Abdul‐Baki, A. A., & Anderson, J. D. (1973). Vigor determination in soybean seed by multiple criteria 1. Crop science, 13(6), 630-633.
Afzal, I., Basra, S. M. A., Ahmad, N., & Farooq, M. (2005). Optimization of hormonal priming techniques for alleviation of salinity stress in wheat (Triticum aestivum L.). Cereal Research Communications, 33(4), 629-634. https://doi.org/10.1007/BF03543587
Aghaei, K., Rah-khosravani, B., Moghanloo, L., & Ghotbi Ravandi, A. A. (2019). Investigating the effect of cadmium accumulation on some biochemical and physiological characteristics in basil (Ocimum basilicum L.). Journal of Plant Process and Function, 8(33), 107-122. [In Persian].
Alharby, H. F., Al-Zahrani, H. S., & Abbas, G. (2022). Potassium and silicon synergistically increase cadmium and lead tolerance and phytostabilization by quinoa through modulation of physiological and biochemical attributes. Toxics, 10(4), 169. https://doi.org/10.3390/toxics10040169
Ameri, A., Fatemi, H., Aroiee, H., & Teixeira da Silva, J.A. (2011). What's the Effect of Saline Priming on Germination Factors of Capsicum annuum var.'California Wonder Seeds? Seed Science and Biotechnology, 5(1), 47-49.
Amini, F., Balouchi, H. R., Movahedi Dehnavi, M., & Attarzadeh, M. (2016). Effect of different concentrations of some heavy metals on germination indices and seed vigor of pinto bean (Phaseolus vulgaris L.). Iranian Journal of Seed Science and Research, 3(2), 95-105. [In Persian].
Amraei, N., & Omidi, H. (2022). Evaluation of the effects of insecticide pre-treatments at different temperatures on germination indices and seedling growth of fuzzy and delinted seeds of two cotton cultivars in the study of seedling growth and germination indices of quinoa genotypes (Chenopodium quinoa Willd.). Seed Research, 12(1), 46-53. [In Persian].
Andrade, V. L., Ribeiro, I., Dos Santos, A. P. M., Aschner, M., & Mateus, M. L. (2023). Metals in Cow Milk and Soy Beverages: Is There a Concern? Toxics, 11(12), 1013. https://doi.org/10.3390/toxics11121013
Ansari, O., Gherekhloo, J., Kamkar, B., and Ghaderi-Far, F. (2016). Breaking seed dormancy and determining cardinal temperatures for Malva sylvestris using nonlinear regression. Seed Science and Technology, 44(3), 1-14.
Apel, K., & Hirt, H. (2004). Reactive oxygen species: metabolism, oxidative stress, and signal transduction. Annual Review of Plant Biology, 55, 373–399. https://doi.org/10.1146/annurev.arplant.55.031903.141701
Arnold, B.R., Fenner, M., & Edwards, P. (1991). Changes in germinability, ABA content, and ABA embryonic sensitivity in developing seeds of Sorghum bicolor (L.) Moench induced by water stress during grain filling. New Phytologist, 118, 339-347.
Atıs, I., Çelik, H., & Ertekin, I. (2024). Germination and early seedling growth in sweet sorghum exposed to heavy metal stress under seed priming pretreatments. International Journal of Agriculture and Wildlife Science, 10(2), 257-272.
Aygun, Y. Z, Atıs, I., &Ertekin, I. (2022). Kadmiyum stresi (CdCl2) altında farklı kinoa (Chenopodium quinoa Willd.) genotiplerinin çimlenme ve ilk fide gelişimi. Mustafa Kemal Üniversitesi Tarım Bilimleri Dergisi, 27(1), 1-8.
Bajji M., Kinet, J.M., & Lutts, S. (2002). Osmotic and ionic effects of NaCl on germination early seedling growth and ion content of Atriplex halimus (chenopodiaceae). Canadian Journal of Botany, 80, 297-304.
Bamagoos, A.A., Alharby, H.F., & Abbas, G. (2022). Differential uptake and translocation of cadmium and lead by quinoa: A multivariate comparison of physiological and oxidative stress responses. Toxics,10(2), 68.
Borsani, O., Valpuesta, V., & Botella, M.A. (2001). Evidence for a role of salicylic acid in the oxidative damage generated by NaCl and osmotic stress in Arabidopsis seedlings. Plant physiology, 126(3), 1024-1030. https://doi.org/10.1104/pp.126.3.1024
Callahan, D. L., Baker, A. J. M., Kolev, S. D., & Wedd, A. G. (2005). Metal ion ligands in hyper accumulating plants. Journal of Biological Inorganic Chemistry, 11(1), 2-12.
Chehregani Rad, A., Farzan, S., & Shirkhani, Z. (2017). A study on the effect of lead treatment on some morphological and physiological indices of petunia (Petunia hybrida L.). Journal of Plant Research (Iranian Journal of Biology), 30(1), 47-57. [In Persian].
Dhindsa, R.S., & Matowe, W. (1981). Drought tolerance in two mosses: correlated with enzymatic defense against lipid peroxidation. Journal of Experimental Botany, 32(1), 79-91.
Dinakar, N., Nagajyoth, P.C., Suresh, S., Damodharam, T., & Suresh, C. (2009). Cadmium induced changes on proline, antioxidant enzymes, nitrate and nitrite reductases in Arachis hypogaea L. Journal of Environmental Biology, 30(2), 289-294.
Esechie, H. A. (1994). Effect of temperature on salinity tolerance of Desmodium at germination stage. In Plant Production on the Threshold of a New Century: Proceedings of the International Conference at the Occasion of the 75th Anniversary of the Wageningen Agricultural University, Wageningen, The Netherlands, held June 28–July 1, 1993 (pp. 447-449). Dordrecht: Springer Netherlands
Gomez, A., Narayan, M., Zhao, L., Jia, X., Bernal, R.A., Lopez-Moreno, M.L., & Peralta-Videa, J.R. (2021). Effects of nano-enabled agricultural strategies on food quality: Current knowledge and future research needs. Journal of Hazardous Materials, 401, 123385. https://doi.org/10.1016/j.jhazmat.2020.123385
Gupta, N., Singh, P.M., Sagar, V., Pandya, A., Chinnappa, M., Kumar, R., & Bahadur, A. (2022). Seed priming with ZnO and Fe3O4 nanoparticles alleviate the Lead toxicity in Basella alba L. through reduced Lead uptake and regulation of ROS. Plants, 11, 2227. https://doi.org/10.3390/plants11172227
Hashemi Shahraki, Sh. (2025). Growth and Biochemical Responses of Sadough Quinoa (Chenopodium quinoa) under Lead Stress. Quality and Durability of Agricultural Products and Food Stuffs, 5(2), 61-71. [In Persian].DOI: https://doi.org/10.71516/qafj.2025.1222724.
Iftikhar, A., Abbas, G., Saqib, M., Shabbir, A., Amjad, M., Shahid, M. (2022). Salinity modulates lead (Pb) tolerance and phytoremediation potential of quinoa: A multivariate comparison of physiological and biochemical attributes. Environmental Geochemistry and Health, 44(1), 257-72.
Ikic, I., Maric evic, M., Tomasovic, S., Gunjaca, J., Atovic, Z. S., & Arcevic, H. S. (2012). The effect of germination temperature on seed dormancy in Croatian-grown winter wheats. Euphytica, 188, 25-34. https://doi.org/10.1007/s10681-012-0695-8
ISTA. (2012). International rules for seed testing, edition 2012. Bassersdorf. Switzerland.
Jacobsen, S.E., Liu, F., & Jensen, C.R. (2009). Does root-sourced ABA play a role for the regulation of stomata under drought in quinoa (Chenopodium quinoa Willd.). Scientia Horticulturae, 122, 281-287.
Jini, D., & Joseph, B. (2017). Physiological mechanism of salicylic acid for alleviation of salt stress in rice. Rice Science, 24(2), 97-108. https://doi.org/10.1016/j.rsci.2016.0
Karim, M.N., Sani, M.N.H., Uddain, J., Azad, M.O.K., Kabir, M.S., Rahman, M.S., Choi, K.Y., & Naznin, T. (2020). Stimulatory effect of seed priming as a pretreatment factor on germination and yield performance of yard long bean (Vigna unguiculata). Horticulturae,6(104), 1-13.
Khan, M.I., Fatma, M., Per T.S., Anjum N., & Khan, A. (2015). Salicylic acid-induced abiotic stress tolerance and underlying mechanisms in plants. Frontiers in Plant Science, 6, 462-770. https://www.ncbi.nlm.nih.gov/pubmed/26175738
Khanna, K., Kohli, S. K., Bali, S., Kaur, P., Saini, P., Bakshi, P., & Bhardwaj, R. (2018). Role of micro-organisms in modulating antioxidant defence in plants exposed to metal toxicity. In plants under metal and metalloid stress, 303 - 335. http://dx.doi.org/10.1016/j.plaphy.2014.07.005
Li, H., Sun, Y. L., Yu, X. H., Guo, H. P., Lian, H. F., Sun, X. D., Shi, Q. H., & Liu, S. Q. (2015). Effects of exogenous calcium on the growth and physiological traits of garlic seedlings under cadmium stress. Journal of Animal and Plant Sciences, 25(3), 107-113. 
Mahdi, I., Fahsi, N., Hafidi, M., Benjelloun, S., Allaoui, A., & Biskri, L. (2021). Rhizospheric phosphate solubilizing Bacillus atrophaeus GQJK17 S8 increases quinoa seedling, withstands heavy metals, and mitigates salt stress. Sustainability, 13(6), 3307. https://doi.org/10.3390/su13063307
Mazhar, M. W., Ishtiaq, M., Maqbool, M., & Akram, R. (2020). Seed priming with Calcium oxide nanoparticles improves germination, biomass, antioxidant defense and yield traits of canola plants under drought stress. South African Journal of Botany, 151, 889-899. https://doi.org/10.1016/j.sajb.2020.10.031
Miller, T., & Chapman, S. J. (1978). Germination responses of three forage grasses to different concentration of six salts. Journal of Range Management, 31(2), 123-124. http://dx.doi.org/10.2307/3897659
Motuzova, G. V., Minkina, T. M., Karpova, E. A., Barsova, N. U., & Mandzhieva, S. S. (2014). Soil contamination with heavy metals as a potential and real risk to the environment. Journal of Geochemical Exploration. 144, 241 - 246. http://dx.doi.org/10.1016/j.jenvman.2013.04.016
Naik, S., Paramesh, R., Siddaraju, R., & Ravi Shankar, P. (2020). Studies on growth parameters in quinoa (Chenopodium quinoa Willd.). International Journal of Chemical Studies, 8(1), 393-397. https://doi.org/10.22271/chemi.2020.v8.i1f.8278
Najafi, Gh., Khomari, S., & Javadi, A. (2015). Germination response of canola seeds to seed vigor changes and hydropriming. Seed Science Research, 45(4), 55-70. https://doi.org/20.1001.1.22520961.1394.5.17.6.9
Nawaz, F., Naeem, M., Akram, A., Ashraf, M.Y., Ahmad, K.S., Zulfiqar, B., Sardar, H., Shabbir, R.N., Majeed, S., Shehzad, M.A., & Anwar, I. (2017). Seed priming with KNO3 mediates biochemical processes to inhibit lead toxicity in maize (Zea mays L.). Journal of the Science of Food and Agriculture, 97(14), 4780-4789.
Nourhosseini, S.A., Safarzadeh, M.N., & Sadeghi, S.M. (2016). Evaluation of germination energy, germination value, and seed vigor indices in peanut (Arachis hypogaea L.). Iranian Journal of Plant Research (Iranian Journal of Biology), 29(1), 221–234. [In Persian]
Nouri, M., & Haddioui, A. (2021). Improving seed germination and seedling growth of Lepidium sativum with different priming methods under arsenic stress. Acta Ecological Sinica, 41(1), 64-71. https://doi.org/10.1016/j.chnaes.2020.08.004
Orchard, T. J. (1977). Estimating the parameters of plant seedling emergence.
Palve, S., Ahire, D., & Gahile, Y. (2022). Salicylic acid pretreatment effects on Beta vulgaris L. multigerm germination and germination indices. International Journal of Biosciences, 20(1), 59-71.
Park, J.H., Choppala, G., Lee, S.J., Bolan, N., Chung, J.W., & Edraki M. (2013). Comparative sorption of Pb and Cd by biochars and its implication for metal immobilization in soils. Water, Air and Soil Pollution, 224: 1-12.
Parmon, G., Ebadi, A., Ghahramani, M., & Mousavi, S. A. (2014). The effect of heavy metals on germination indices and vigor of maize seeds under laboratory conditions. Seed Research, 4(12), 40-51. [In Persian].
Peralta, J.R., Gardea-Torresdey, J.L., & Tiemann, K.J. (2000). Study of the effects of heavy metals on seed germination and plant growth on alfalfa plant (Medicago sativa) grown in solid media. Proceedings of the 2000 Conference on Hazardous Waste Research, pp. 135-140.
Pourrut, B., Shahid, M., Dumat, C., Winterton, P., & Pinelli, E. (2011). Lead uptake, toxicity, and detoxification in plants. Reviews of Environmental Contamination and Toxicology, 213, 113 - 136. http://dx.doi.org/10.1016/j.jhazmat.2010.10.047
Prasad, A.S. (2003). Zinc deficiency. British Medical Journal. 326: 409-410.
Rahman Khan, M., & Mahmud Khan, M. (2010). Effect of varying concentrations of nickel and cobalt on the plant growth and yield of chickpea. Australian Journal Basic and Applied Science, 4(6), 1036-104.
Rezaei, B., Amirinejad, A. A., & Ghobadi, M. (2022). Interaction effects of lead nitrate, salicylic acid, and biochar on the growth characteristics of thyme (Thymus vulgaris L.). Journal of Water and Soil, 36(1), 67-79. [In Persian].
Rizwan, M., Ali, S., Ali, B., Adrees, M., Arshad, M., Hussain, A., & Waris, A. A. (2019). Zinc and iron oxide nanoparticles improved the plant growth and reduced the oxidative stress and cadmium concentration in wheat. Chemosphere, 214, 269-277. https://doi.org/10.1016/j.chemosphere.2018.09.120
Roosta, H.R., Jalali, M., & Ali Vakili Shahrbabaki, S.M. (2015). Effect of Nano Fe-Chelate, FeEddha and FeSO4on Vegetative Growth, Physiological Parameters and Some Nutrient Elements Concentrations of Four Varieties of Lettuce (Lactuca sativa L.) In NFT System. Jornal of Plant Nutrtent, 38, 2176–2184. https://doi.org/10.1080/01904167.2015.1043378
Sekabira, K., Oryem- Origa, H., Mutumba, G., Kakudidi, E., & Basamba, T.A. (2011). Heavy metal phytoremediation by Commelina benghalensis (L) and Cynodon dactylon (L) growing in Urban stream sediments. International Journal of Plant Physiology and Biochemistry, 3(8), 133-142.
Shah, A. A., Ahmed, S., Abbas, M., & Yasin, N. A. (2020). Seed priming with 3-epibrassinolide alleviates cadmium stress in Cucumis sativus through modulation of antioxi-dative system and gene expression. Scientia Horticulturae, 265, 109203. https://doi.org/10.1016/j.scienta.2020.109203
Soltanzadeh Pormehr, S., Maleki Lajayer, H., Torabi Giglou, M., Esmaielpour, B., Chamani, E., & Pourbeyrami Hir, Y. (2021). Effects of dill seed priming with salicylic acid under lead and cadmium heavy metal stress. Journal of Seed Research, 4(3), 33-44. [In Persian].
Taie, H. A., El-Yazal, M. A. S., Ahmed, S. M., & Rady, M. M. (2019). Polyamines modulate growth, antioxidant activity, and genomic DNA in heavy metal-stressed wheat plant. Environmental Science and Pollution Research, 26, 22338-22350.
Vahdani Rashvanlouei, M., Jami Al-Ahmadi, M., Siari Zehan, M.H., Shorideh, H., & Mostafaei, M. (2023). Effect of seed nutritional priming with iron sulfate and zinc sulfate on germination characteristics and seedling growth of lentil seeds. Seed Research, 13(2), 1–14. [In Persian]
Venkatachalam, P., Jayalakshmi, N., Geetha, N., Sahi, S. V., Sharma, N. C., Rene, E. R., Sarkar, S. K., & Favas P. J. C. (2017) Accumulation efficiency, genotoxicity and antioxidant defense mechanisms in medicinal plant Acalypha indica L. under lead stress. Chemosphere, 171, 544-553.
Wang, W., He, A., Peng, S., Huang, J., Cui, K., & Nie, L. (2018). The effect of storage condition and duration on the deterioration of primed rice seeds. Frontiers in Plant Science, 9, 1-17.
Xin, Y., Liu, M., Wei, L., Gao, Y., Ruan, Y., Wang, Q., & Zhang, Z. (2023). Changes in soil chemical properties and rhizosphere bacterial community induced by soil amendments associated with reduction in cadmium accumulation by rice. Agronomy, 13(12), 3051. https://doi.org/10.3390/agronomy13123051
Yang, Y., Wei, X., Lu, J., You, J., Wang, W., & Shi, R. (2010). Lead-induced phytotoxicity mechanism involved in seed germination and seedling growth of wheat (Triticum aestivum L.). Ecotoxicology and Environmental Safety, 73(8): 1982-1987.
Yang, Y., Zhang, Y., Wei, X., You, J., Wang, W., Lu, J., & Shi, R. (2011). Comparative antioxidative responses and proline metabolism in two wheat cultivars under short term lead stress. Ecotoxicology and Environmental Safety 74, 733-740.
Zhou, P., Adeel, M., Shakoor, N., Guo, M., Hao, Y., & Azeem, I. (2021). Application of nanoparticles alleviates heavy metals stress and promotes plant growth: An overview. Nanomaterials, 11, 26-38.