{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,8]],"date-time":"2026-09-08T11:07:27Z","timestamp":1788865647833,"version":"build-2803163510"},"reference-count":102,"publisher":"MDPI AG","issue":"18","license":[{"start":{"date-parts":[[2026,9,8]],"date-time":"2026-09-08T00:00:00Z","timestamp":1788825600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"REASILIENCE Project","award":["SBEP571"],"award-info":[{"award-number":["SBEP571"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>Halophytes are recognized for their adaptive capacity and potential applications in phytomanagement and ecosystem restoration. This study investigates the comparative physiology and antioxidant responses of two native halophytes, Caroxylon vermiculatum (L.) and Mesembryanthemum nodiflorum L., growing naturally in saline and metal-affected coastal environments. A comprehensive set of physiological and biochemical parameters was assessed, including macro- and microelements (Na, K, Ca, Fe, Zn, Cd), photosynthetic pigments (chlorophyll a, chlorophyll b, and carotenoids), soluble sugars, and proteins. In addition, oxidative stress markers (hydrogen peroxide, H2O2, and malondialdehyde, MDA), enzymatic antioxidants (superoxide dismutase, SOD, catalase, CAT, and guaiacol peroxidase, GPX), and non-enzymatic antioxidants (total phenolics, flavonoids, and proanthocyanidins) were evaluated. Antioxidant activities, including DPPH (2,2-diphenyl-1-picrylhydrazyl) radical scavenging and reducing power, were also measured. The results revealed clear species-specific adaptive strategies. Mesembryanthemum nodiflorum exhibited higher accumulation of Na and K, together with elevated levels of carotenoids and oxidative stress markers. This species showed translocation factors (TF &gt; 1) for Na (~1.60) and K (~1.90), indicating efficient ion transport to aerial parts, while displaying low bioconcentration (BCF &lt; 0.5 for most trace elements) and biological accumulation factors (BAF &lt; 1), suggesting limited capacity for heavy metal accumulation. In contrast, Caroxylon vermiculatum showed higher concentrations of chlorophylls, carotenoids, phenolic compounds, flavonoids, and proteins, along with stronger superoxide dismutase activity. It also exhibited lower translocation of trace elements (TF &lt; 1) and higher root retention of metals (BCF up to ~0.5), indicating a more effective exclusion and detoxification strategy. Overall, these findings demonstrate that M. nodiflorum relies on ion accumulation and translocation, whereas C. vermiculatum exhibits stronger ion regulation and antioxidant protection. Given the moderate BAF and BCF values observed, both species are more likely to contribute to phytomanagement through ion regulation and phytostabilization rather than efficient phytoextraction.<\/jats:p>","DOI":"10.3390\/plants15182743","type":"journal-article","created":{"date-parts":[[2026,9,8]],"date-time":"2026-09-08T10:00:04Z","timestamp":1788861604000},"page":"2743","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Mineral Accumulation and Physiological Responses of Two Halophytes, Caroxylon vermiculatum and Mesembryanthemum nodiflorum: Implications for Phytoremediation in Contaminated Coastal Environments"],"prefix":"10.3390","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-5733-8377","authenticated-orcid":false,"given":"Dhouha Belhadj","family":"Sghaier","sequence":"first","affiliation":[{"name":"National Institute of Marine Sciences and Technologies, University of Carthage, Tunis 2025, Tunisia"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"conceptualization","vocabulary":"credit"},{"role":"methodology","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"},{"role":"formal-analysis","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9575-4320","authenticated-orcid":false,"given":"Hasna","family":"Ellouzi","sequence":"additional","affiliation":[{"name":"Laboratory of Extremophile Plants, Centre of Biotechnology of Borj Cedria, Hammam-Lif 2050, Tunisia"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"methodology","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"given":"Houyem","family":"Abderrazak","sequence":"additional","affiliation":[{"name":"National Institute of Research and Physicochemical Analysis, Sidi Thabet Technopole, Ariana 2020, Tunisia"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"methodology","vocabulary":"credit"},{"role":"formal-analysis","vocabulary":"credit"}]},{"given":"Fourat","family":"Akrout","sequence":"additional","affiliation":[{"name":"National Institute of Marine Sciences and Technologies, University of Carthage, Tunis 2025, Tunisia"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"methodology","vocabulary":"credit"},{"role":"formal-analysis","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4409-2755","authenticated-orcid":false,"given":"Mohsen","family":"Hanana","sequence":"additional","affiliation":[{"name":"Laboratory of Extremophile Plants, Centre of Biotechnology of Borj Cedria, Hammam-Lif 2050, Tunisia"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"writing-review-editing","vocabulary":"credit"},{"role":"supervision","vocabulary":"credit"},{"role":"project-administration","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0651-1605","authenticated-orcid":false,"given":"Monia EL","family":"Bour","sequence":"additional","affiliation":[{"name":"National Institute of Marine Sciences and Technologies, University of Carthage, Tunis 2025, Tunisia"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"writing-review-editing","vocabulary":"credit"},{"role":"supervision","vocabulary":"credit"},{"role":"project-administration","vocabulary":"credit"}]}],"member":"1968","published-online":{"date-parts":[[2026,9,8]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"327","DOI":"10.1093\/aob\/mcu267","article-title":"Plant salt tolerance: Adaptations in halophytes","volume":"115","author":"Flowers","year":"2015","journal-title":"Ann. Bot."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"104601","DOI":"10.1016\/j.envexpbot.2021.104601","article-title":"Halophyte based Mediterranean agriculture in the contexts of food insecurity and global climate change","volume":"191","author":"Castagna","year":"2021","journal-title":"Environ. Exp. Bot."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"651","DOI":"10.1146\/annurev.arplant.59.032607.092911","article-title":"Mechanisms of salinity tolerance","volume":"59","author":"Munns","year":"2008","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_4","unstructured":"Food and Agriculture Organization of the United Nations (FAO) (2024). Global Status of Salt-Affected Soils, FAO."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"1707","DOI":"10.1016\/j.biochi.2006.07.003","article-title":"Toxic metal accumulation, responses to exposure and mechanisms of tolerance in plants","volume":"88","author":"Clemens","year":"2006","journal-title":"Biochimie"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1093\/jxb\/erq281","article-title":"Root responses to cadmium in plants","volume":"62","author":"Lux","year":"2011","journal-title":"J. Exp. Bot."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Kumar, P., and Sharma, P.K. (2020). Soil Salinity and Food Security in India. Front. Sustain. Food Syst., 4.","DOI":"10.3389\/fsufs.2020.533781"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"33017","DOI":"10.1073\/pnas.2013771117","article-title":"Predicting long-term dynamics of soil salinity and sodicity on a global scale","volume":"117","author":"Hassani","year":"2020","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Rashid, A., Schutte, B.J., Ulery, A., Deyholos, M.K., Sanogo, S., Lehnhoff, E.A., and Beck, L. (2023). Heavy Metal Contamination in Agricultural Soil: Environmental Pollutants Affecting Crop Health. Agronomy, 13.","DOI":"10.20944\/preprints202305.0398.v1"},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Yan, A., Wang, Y., Tan, S.N., Yusof, L.K.M., Ghosh, S., and Chen, Z. (2020). Phytoremediation: A Promising Approach for Revegetation of Heavy Metal-Polluted Land. Front. Plant Sci., 11.","DOI":"10.3389\/fpls.2020.00359"},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Alengebawy, A., Abdelkhalek, S.T., Qureshi, S.R., and Wang, M.Q. (2021). Heavy Metals and Pesticides Toxicity in Agricultural Soil and Plants: Ecological Risks and Human Health Implications. Toxics, 9.","DOI":"10.3390\/toxics9030042"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"11786302231201259","DOI":"10.1177\/11786302231201259","article-title":"Heavy Metal Pollution in the Environment and Its Impact on Health: Exploring Green Technology for Remediation","volume":"17","author":"Das","year":"2023","journal-title":"Environ. Health Insights"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"434","DOI":"10.1016\/j.ecofro.2025.10.009","article-title":"A comprehensive review on potential heavy metals in the environment: Persistence, bioaccumulation, ecotoxicology, and agricultural impacts","volume":"46","author":"Mustapha","year":"2025","journal-title":"Ecol. Front."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"134788","DOI":"10.1016\/j.chemosphere.2022.134788","article-title":"Phytoremediation of heavy metals in soil and water: An eco-friendly, sustainable and multidisciplinary approach","volume":"301","author":"Bhat","year":"2022","journal-title":"Chemosphere"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1016\/j.bcab.2017.11.003","article-title":"Differential behavior of the antioxidant system in response to salinity induced oxidative stress in salt-tolerant and salt-sensitive cultivars of Brassica juncea L.","volume":"13","author":"Kumar","year":"2017","journal-title":"Biocatal. Agric. Biotechnol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"56","DOI":"10.1038\/s44264-025-00098-6","article-title":"Agricultural practices can threaten soil resilience through changing feedback loops","volume":"3","author":"Carswell","year":"2025","journal-title":"npj Sustain. Agric."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"1805902","DOI":"10.1080\/15592324.2020.1805902","article-title":"Heavy metal tolerance and potential for remediation of heavy metal-contaminated saline soils for the euhalophyte Suaeda salsa","volume":"15","author":"Shang","year":"2020","journal-title":"Plant Signal. Behav."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"105397","DOI":"10.1016\/j.envexpbot.2023.105397","article-title":"Sustainable agricultural management of saline soils in arid and semi-arid Mediterranean regions through halophytes, microbial and soil-based technologies","volume":"212","author":"Nogales","year":"2023","journal-title":"Environ. Exp. Bot."},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"Hasanuzzaman, M., Bhuyan, M.H.M.B., Zulfiqar, F., Raza, A., Mohsin, S.M., Mahmud, J.A., Fujita, M., and Fotopoulos, V. (2020). Reactive Oxygen Species and Antioxidant Defense in Plants under Abiotic Stress: Revisiting the Crucial Role of a Universal Defense Regulator. Antioxidants, 9.","DOI":"10.3390\/antiox9080681"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"33","DOI":"10.1016\/j.tplants.2014.07.005","article-title":"Iron nutrition, signaling, and homeostasis in plants","volume":"20","author":"Briat","year":"2015","journal-title":"Trends Plant Sci."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"100057","DOI":"10.1016\/j.ncrops.2024.100057","article-title":"Genomic and modern biotechnological strategies for enhancing salt tolerance in crops","volume":"2","author":"Yuan","year":"2025","journal-title":"New Crops"},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Sharma, J.K., Kumar, N., Singh, N.P., and Santal, A.R. (2023). Phytoremediation technologies and their mechanism for removal of heavy metal from contaminated soil: An approach for a sustainable environment. Front. Plant Sci., 14.","DOI":"10.3389\/fpls.2023.1076876"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"137524","DOI":"10.1016\/j.chemosphere.2022.137524","article-title":"Halophytes for the sustainable remediation of heavy metal-contaminated sites: Recent developments and future perspectives","volume":"313","author":"Singh","year":"2023","journal-title":"Chemosphere"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"105797","DOI":"10.1016\/j.ecoleng.2020.105797","article-title":"Phyto-extraction of zinc, lead, nickel, and cadmium from zinc leach residue by a halophyte: Salicornia europaea","volume":"148","author":"Khanlarian","year":"2020","journal-title":"Ecol. Eng."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"123368","DOI":"10.1016\/j.jhazmat.2020.123368","article-title":"Salinity alleviates the arsenic toxicity in the facultative halophyte Salvadora persica L. by the modulations of physiological, biochemical, and ROS scavenging attributes","volume":"401","author":"Patel","year":"2021","journal-title":"J. Hazard. Mater."},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Ibraheem, F., Al-Zahrani, A., and Mosa, A. (2022). Physiological Adaptation of Three Wild Halophytic Suaeda Species: Salt Tolerance Strategies and Metal Accumulation Capacity. Plants, 11.","DOI":"10.3390\/plants11040537"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"323","DOI":"10.1007\/s11104-019-04152-4","article-title":"Halophyte Halogeton glomeratus, a promising candidate for phytoremediation of heavy metal-contaminated saline soils","volume":"442","author":"Li","year":"2019","journal-title":"Plant Soil"},{"key":"ref_28","first-page":"301","article-title":"Assessment of phytoremediation capacity of three halophytes: Suaedamonoica, Tamarix indica and Cressa critica","volume":"71","author":"Joshi","year":"2020","journal-title":"BiologiaFutur."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"72","DOI":"10.1007\/s41742-025-00738-2","article-title":"Screening for Heavy Metal-Resistant Clones in the Xero-Halophyte Atriplex halimus L.: A Prerequisite for Phytoremediation of Polymetallic Mining Pollution in Arid Areas","volume":"19","author":"Zaghdoudi","year":"2025","journal-title":"Int. J. Environ. Res."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"161","DOI":"10.23902\/trkjnat.1458374","article-title":"A new species record for the flora of T\u00fcrkiye; Caroxylon vermiculatum (L.) Akhani&Roalson (Chenopodiaceae\/Amaranthaceae)","volume":"25","author":"Yaprak","year":"2024","journal-title":"Trak. Univ. J. Nat. Sci."},{"key":"ref_31","doi-asserted-by":"crossref","unstructured":"Lima, A.R., Gama, F., Casta\u00f1eda-Loaiza, V., Costa, C., Sch\u00fcler, L.M., Santos, T., Salazar, M., Nunes, C., Cruz, R.M.S., and Varela, J. (2021). Nutritional and Functional Evaluation of Inula crithmoides and Mesembryanthemum nodiflorum Grown in Different Salinities for Human Consumption. Molecules, 26.","DOI":"10.3390\/molecules26154543"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"509","DOI":"10.1093\/aob\/mcu264","article-title":"How can we take advantage of halophyte properties to cope with heavy metal toxicity in salt-affected areas?","volume":"115","author":"Lutts","year":"2015","journal-title":"Ann. Bot."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"269","DOI":"10.1139\/er-2016-0063","article-title":"Phytoremediation of heavy metal contaminated saline soils using halophytes: Current progress and future perspectives","volume":"25","author":"Liang","year":"2017","journal-title":"Environ. Rev."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"134235","DOI":"10.1016\/j.scitotenv.2019.134235","article-title":"Curing the earth: A review of anthropogenic soil salinization and plant-based strategies for sustainable mitigation","volume":"698","author":"Litalien","year":"2020","journal-title":"Sci. Total Environ."},{"key":"ref_35","doi-asserted-by":"crossref","unstructured":"Hasanuzzaman, M., and Fujita, M. (2023). Plant Responses and Tolerance to Salt Stress: Physiological and Molecular Interventions. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms242115740"},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Isayenkov, S.V., and Maathuis, F.J.M. (2019). Plant salinity stress: Many unanswered questions remain. Front. Plant Sci., 10.","DOI":"10.3389\/fpls.2019.00080"},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Zhan, N., Yang, H., Li, J., Shi, X., Yang, B., Sun, Y., Guo, G., and Cui, X. (2025). Spatial Distribution Characteristics and Relationships of Salt-Based Ions and Nutrients in Old Protected Vegetable Fields. Horticulturae, 11.","DOI":"10.3390\/horticulturae11020126"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"172564","DOI":"10.1016\/j.scitotenv.2024.172564","article-title":"Nutrient dynamics in wetland systems associated with hydrological and anthropogenic variations in the south of Samboromb\u00f3n Bay, Argentina","volume":"928","author":"Carol","year":"2024","journal-title":"Sci. Total Environ."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"de Vicente, I. (2021). Biogeochemistry of Mediterranean Wetlands: A Review about the Effects of Water-Level Fluctuations on Phosphorus Cycling and Greenhouse Gas Emissions. Water, 13.","DOI":"10.3390\/w13111510"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"578","DOI":"10.1111\/ppl.13185","article-title":"Ion homeostasis for salinity tolerance in plants: A molecular approach","volume":"171","author":"Amin","year":"2021","journal-title":"Physiol. Plant."},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Mansoor, S., Ali, A., Kour, N., Bornhorst, J., AlHarbi, K., Rinklebe, J., Abd El Moneim, D., Ahmad, P., and Chung, Y.S. (2023). Heavy Metal Induced Oxidative Stress Mitigation and ROS Scavenging in Plants. Plants, 12.","DOI":"10.3390\/plants12163003"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"124922","DOI":"10.1016\/j.chemosphere.2019.124922","article-title":"Soil organic matter prevails over heavy metal pollution and vegetation as a factor shaping soil microbial communities at historical Zn-Pb mining sites","volume":"240","author":"Stefanowicz","year":"2020","journal-title":"Chemosphere"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"1032","DOI":"10.1111\/plb.12989","article-title":"The speciation and distribution characteristics of Cu in Phragmites australis (Cav.) Trin ex. Steudel","volume":"21","author":"Wu","year":"2019","journal-title":"Plant Biol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"104567","DOI":"10.1016\/j.bse.2022.104567","article-title":"Evaluation of osmoregulation and morpho-physiological responses of Borago officinalis under drought and salinity stress with equal osmotic potential","volume":"106","author":"Rostampour","year":"2023","journal-title":"Biochem. Syst. Ecol."},{"key":"ref_45","doi-asserted-by":"crossref","unstructured":"Hashemipetroudi, S., Ahmadian, G., Fatemi, F., Nematzadeh, G., Yamchi, A., and Kuhlmann, M. (2022). Ion content, antioxidant enzyme activity and transcriptional response under salt stress and recovery condition in the halophyte grass Aeluropus littoralis. BMC Res. Notes, 15.","DOI":"10.1186\/s13104-022-06090-4"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"2112","DOI":"10.1111\/ppl.13446","article-title":"Seasonal environmental changes affect differently the physiological and biochemical responses of two Limonium species in Sabkha biotope","volume":"172","author":"Maaloul","year":"2021","journal-title":"Physiol. Plant."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"983","DOI":"10.1111\/pbi.14553","article-title":"The natural variation in shoot Na+ content and salt tolerance in maize is attributed to various minor-effect variants, including an SNP located in the promoter of ZmHAK11","volume":"23","author":"Liang","year":"2025","journal-title":"Plant Biotechnol. J."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"738","DOI":"10.1111\/1440-1703.12345","article-title":"Physiological responses of the xero-halophyte Salsola drummondii to seasonal alterations of environmental conditions in a salt desert","volume":"37","author":"Rasheed","year":"2022","journal-title":"Ecol. Res."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"204","DOI":"10.1016\/j.jphotobiol.2015.06.003","article-title":"Growth, chlorophyll fluorescence and mineral nutrition in the halophyte Tamarix gallica cultivated in combined stress conditions: Arsenic and NaCl","volume":"149","author":"Sghaier","year":"2015","journal-title":"J. Photochem. Photobiol. B Biol."},{"key":"ref_50","first-page":"153","article-title":"Ionic homeostasis and oxidative stress tolerance in plants","volume":"37","author":"Bose","year":"2014","journal-title":"Plant Cell Environ."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1016\/j.cj.2018.01.003","article-title":"Plant salt tolerance and Na+ sensing and transport","volume":"6","author":"Wu","year":"2018","journal-title":"Crop J."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"949","DOI":"10.1071\/FP16007","article-title":"Stress tolerance mechanisms in Juncus: Responses to salinity and drought in three Juncus species adapted to different natural environments","volume":"43","author":"Hassan","year":"2016","journal-title":"Funct. Plant Biol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"100034","DOI":"10.1016\/j.ncrops.2024.100034","article-title":"Research progress on the physiological and molecular mechanisms underlying soybean aluminum resistance","volume":"2","author":"Li","year":"2025","journal-title":"New Crops"},{"key":"ref_54","doi-asserted-by":"crossref","unstructured":"Rahman, M., Mostofa, M., Keya, S., Siddiqui, M., Ansary, M., Das, A., Rahman, M., and Tran, L. (2021). Adaptive Mechanisms of Halophytes and Their Potential in Improving Salinity Tolerance in Plants. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms221910733"},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Ltaeif, H., Sakhraoui, A., Gonz\u00e1lez-Orenga, S., Faz, A., Boscaiu, M., Vicente, \u00d3., and Rouz, S. (2021). Responses to Salinity in Four Plantago Species from Tunisia. Plants, 10.","DOI":"10.3390\/plants10071392"},{"key":"ref_56","doi-asserted-by":"crossref","unstructured":"Azizi, M., Faz, A., Zornoza, R., Martinez-Martinez, S., and Acosta, J.A. (2023). Phytoremediation Potential of Native Plant Species in Mine Soils Polluted by Metal(loid)s and Rare Earth Elements. Plants, 12.","DOI":"10.3390\/plants12061219"},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Ioni\u021b\u0103, M.F., Dunca, E.C., and Radu, S.M. (2026). Molecular and Cellular Mechanisms of Plant Responses to Heavy Metal Stress in Mining-Impacted Environments. Plants, 15.","DOI":"10.3390\/plants15071045"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"102","DOI":"10.1007\/s11738-016-2113-y","article-title":"Chlorophyll a fluorescence as a tool to monitor physiological status of plants under abiotic stress conditions","volume":"38","author":"Kalaji","year":"2016","journal-title":"Acta Physiol. Plant."},{"key":"ref_59","doi-asserted-by":"crossref","unstructured":"Sasi, S., Venkatesh, J., Daneshi, R., and Gururani, M. (2018). Photosystem II Extrinsic Proteins and Their Putative Role in Abiotic Stress Tolerance in Higher Plants. Plants, 7.","DOI":"10.3390\/plants7040100"},{"key":"ref_60","first-page":"806","article-title":"Sorghum and salinity: II. Gas exchange and chlorophyll fluorescence of sorghum under salt stress","volume":"44","author":"Netondo","year":"2004","journal-title":"Crop Sci."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"306","DOI":"10.1016\/j.sajb.2016.03.011","article-title":"Effect of salt stress on growth, chlorophyll content, lipid peroxidation and antioxidant defence systems in Phaseolus vulgaris L.","volume":"105","author":"Abderrahim","year":"2016","journal-title":"S. Afr. J. Bot."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"597","DOI":"10.1111\/tpj.12386","article-title":"Carotenoid oxidation products as stress signals in plants","volume":"79","author":"Havaux","year":"2014","journal-title":"Plant J."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"21512","DOI":"10.1038\/s41598-024-72627-1","article-title":"Carbohydrate accumulation patterns in mangrove and halophytic plant species under seasonal variation","volume":"14","author":"Mehta","year":"2024","journal-title":"Sci. Rep."},{"key":"ref_64","doi-asserted-by":"crossref","unstructured":"Rao, M.J., Duan, M., Ikram, M., and Zheng, B. (2025). ROS Regulation and Antioxidant Responses in Plants Under Air Pollution: Molecular Signaling, Metabolic Adaptation, and Biotechnological Solutions. Antioxidants, 14.","DOI":"10.3390\/antiox14080907"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"1242","DOI":"10.1111\/pce.12061","article-title":"Plant sugars are crucial players in the oxidative challenge during abiotic stress: Extending the traditional concept","volume":"36","author":"Keunen","year":"2013","journal-title":"Plant Cell Environ."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"9819314","DOI":"10.34133\/2022\/9819314","article-title":"Biological Parts for Engineering Abiotic Stress Tolerance in Plants","volume":"2022","author":"Lohani","year":"2022","journal-title":"Biodes. Res."},{"key":"ref_67","first-page":"1756","article-title":"Sensing \u03b2-carotene oxidation in photosystem II to master plant stress tolerance","volume":"223","author":"Havaux","year":"2019","journal-title":"New Phytol."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"1159","DOI":"10.3390\/molecules17021159","article-title":"Primary, secondary metabolites, H2O2, malondialdehyde and photosynthetic responses of Orthosiphon stimaneus Benth. to different irradiance levels","volume":"17","author":"Ibrahim","year":"2012","journal-title":"Molecules"},{"key":"ref_69","doi-asserted-by":"crossref","unstructured":"Mishra, N., Jiang, C., Chen, L., Paul, A., Chatterjee, A., and Shen, G. (2023). Achieving abiotic stress tolerance in plants through antioxidative defense mechanisms. Front. Plant Sci., 14.","DOI":"10.3389\/fpls.2023.1110622"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"779","DOI":"10.1007\/s10265-021-01285-5","article-title":"Salinity induced alterations in photosynthetic and oxidative regulation are ameliorated as a function of salt secretion","volume":"134","author":"Hussain","year":"2021","journal-title":"J. Plant Res."},{"key":"ref_71","doi-asserted-by":"crossref","unstructured":"Ievina, S., Karlsons, A., Osvalde, A., Andersone-Ozola, U., and Ievins, G. (2023). Coastal Wetland Species Rumex hydrolapathum: Tolerance against Flooding, Salinity, and Heavy Metals for Its Potential Use in Phytoremediation and Environmental Restoration Technologies. Life, 13.","DOI":"10.3390\/life13071604"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"112368","DOI":"10.1016\/j.ecoenv.2021.112368","article-title":"Evaluation of phytoremediation potential of native dominant plants and spatial distribution of heavy metals in abandoned mining area in Southwest China","volume":"220","author":"Wu","year":"2021","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"ref_73","doi-asserted-by":"crossref","unstructured":"Zandi, P., and Schnug, E. (2022). Reactive Oxygen Species, Antioxidant Responses and Implications from a Microbial Modulation Perspective. Biology, 11.","DOI":"10.3390\/biology11020155"},{"key":"ref_74","doi-asserted-by":"crossref","unstructured":"Mujeeb, A., Abideen, Z., Aziz, I., Sharif, N., Hussain, M.I., Qureshi, A.S., and Yang, H.H. (2023). Phytoremediation of Potentially Toxic Elements from Contaminated Saline Soils Using Salvadora persica L.: Seasonal Evaluation. Plants, 12.","DOI":"10.3390\/plants12030598"},{"key":"ref_75","doi-asserted-by":"crossref","unstructured":"Pirasteh-Anosheh, H., Samadi, M., Kazemeini, S.A., Ozturk, M., Ludwiczak, A., and Piernik, A. (2023). ROS Homeostasis and Antioxidants in the Halophytic Plants and Seeds. Plants, 12.","DOI":"10.3390\/plants12173023"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"100005","DOI":"10.1016\/j.ncrops.2023.11.003","article-title":"The central role of transcription factors in bridging biotic and abiotic stress responses for plants\u2019 resilience","volume":"1","author":"Liu","year":"2024","journal-title":"New Crops"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"570","DOI":"10.3390\/stresses3030040","article-title":"An Overview of Plant Phenolics and Their Involvement in Abiotic Stress Tolerance","volume":"3","author":"Kumar","year":"2023","journal-title":"Stresses"},{"key":"ref_78","doi-asserted-by":"crossref","unstructured":"Rao, M., and Zheng, B. (2025). The Role of Polyphenols in Abiotic Stress Tolerance and Their Antioxidant Properties to Scavenge Reactive Oxygen Species and Free Radicals. Antioxidants, 14.","DOI":"10.3390\/antiox14010074"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"27","DOI":"10.62425\/crihs.1628360","article-title":"Antioxidant Potential and Phenolic Content of Plantago major","volume":"2","author":"Atmaca","year":"2025","journal-title":"Curr. Health Sci. J."},{"key":"ref_80","doi-asserted-by":"crossref","unstructured":"Souza, M., Silva, B., Badiale-Furlong, E., and Costa, C. (2021). Phenolic Acid Profile, Quercetin Content, and Antioxidant Activity of Six Brazilian Halophytes. Handbook of Halophytes, Springer.","DOI":"10.1007\/978-3-030-57635-6_44"},{"key":"ref_81","doi-asserted-by":"crossref","unstructured":"Brzovi\u0107, P., Radman, S., and Mekini\u0107, I.G. (2025). Phenolic Antioxidants in the Adriatic Halophyte Limbardacrithmoides: Variation Across Phenological Stages. Foods, 14.","DOI":"10.3390\/foods14213718"},{"key":"ref_82","doi-asserted-by":"crossref","unstructured":"Ja\u0144czak-Pieni\u0105\u017cek, M., Cicho\u0144ski, J., Michalik, P., and Chrzanowski, G. (2022). Effect of Heavy Metal Stress on Phenolic Compounds Accumulation in Winter Wheat Plants. Molecules, 28.","DOI":"10.3390\/molecules28010241"},{"key":"ref_83","doi-asserted-by":"crossref","unstructured":"Jarin, A.S., Khan, M.A.R., Apon, T.A., Islam, M.A., Rahat, A., Akter, M., Anik, T.R., Nguyen, H.M., Nguyen, T.T., and Ha, C.V. (2025). Plant Responses to Heavy Metal Stresses: Mechanisms, Defense Strategies, and Nanoparticle-Assisted Remediation. Plants, 14.","DOI":"10.3390\/plants14243834"},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"244","DOI":"10.1016\/j.plaphy.2007.02.001","article-title":"Salinity effects on polyphenol content and antioxidant activities in leaves of the halophyte Cakile maritima","volume":"45","author":"Ksouri","year":"2007","journal-title":"Plant Physiol. Biochem."},{"key":"ref_85","first-page":"1","article-title":"Land use and recent morphodynamics in the Ariana River sebkha watershed (North-eastern Tunisia)","volume":"24","author":"Chouari","year":"2015","journal-title":"Confins"},{"key":"ref_86","unstructured":"Chouari, W. (2009). Environment and Natural Risks in Greater Tunis, Cartographic Approach. [Ph.D. Thesis, University of Paris VII]."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"716","DOI":"10.1007\/s12517-022-09877-8","article-title":"Health risk assessment associated to heavy metal pollution levels in Mediterranean environment soils: A case study in the watershed of Sebkhet Ariana, Tunisia","volume":"15","author":"Ghouma","year":"2022","journal-title":"Arab. J. Geosci."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"110360","DOI":"10.1016\/j.scienta.2021.110360","article-title":"Seed-priming with H2O2 alleviates subsequent salt stress by preventing ROS production and amplifying antioxidant defense in cauliflower seeds and seedlings","volume":"288","author":"Ellouzi","year":"2021","journal-title":"Sci. Hortic."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"13063","DOI":"10.15835\/nbha51213063","article-title":"Phytoremediation potential of native plants: Biomonitoring approach in contaminated soils approach in contaminated soils","volume":"51","author":"Aydi","year":"2023","journal-title":"Not. Bot. Horti Agrobot."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"497","DOI":"10.1007\/BF03325986","article-title":"Speciation of heavy metals in electroplating industry sludge and wastewater residue using inductively coupled plasma","volume":"4","author":"Venkateswaran","year":"2007","journal-title":"Int. J. Environ. Sci. Technol."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"87","DOI":"10.1080\/15226514.2016.1216078","article-title":"Phytoremediation of industrial mines wastewater using water hyacinth","volume":"19","author":"Saha","year":"2017","journal-title":"Int. J. Phytoremediat."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"350","DOI":"10.1016\/0076-6879(87)48036-1","article-title":"Chlorophylls and carotenoids: Pigments of photosynthetic biomembranes","volume":"148","author":"Lichtenthaler","year":"1987","journal-title":"Methods Enzymol."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"604","DOI":"10.1007\/s004250050524","article-title":"Improving the thiobarbituric acid-reactive-substances assay for estimating lipid peroxidation in plant tissues containing anthocyanin and other interfering compounds","volume":"207","author":"Hodges","year":"1999","journal-title":"Planta"},{"key":"ref_94","first-page":"121","article-title":"Effect of spermine, atrazine and combination between them on some endogenous protective systems and stress markers in plants","volume":"51","author":"Sergiev","year":"1997","journal-title":"Compt. Rend. Acad. Bulg. Sci."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"2090","DOI":"10.1248\/cpb.36.2090","article-title":"Two new flavonoids and other constituents in licorice root: Their relative astringency and radical scavenging effects","volume":"36","author":"Hatano","year":"1988","journal-title":"Chem. Pharm. Bull."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"248","DOI":"10.1016\/0003-2697(76)90527-3","article-title":"A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding","volume":"72","author":"Bradford","year":"1976","journal-title":"Anal. Biochem."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"307","DOI":"10.5264\/eiyogakuzashi.44.307","article-title":"Studies on products of browning reaction","volume":"44","author":"Oyaizu","year":"1986","journal-title":"Jpn. J. Soc. Nutr. Diet."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"38","DOI":"10.1016\/j.jplph.2016.11.014","article-title":"H2O2 seed priming improves tolerance to salinity, drought and their combined effect more than mannitol in Cakile maritima when compared to Eutrema salsugineum","volume":"210","author":"Ellouzi","year":"2017","journal-title":"J. Plant Physiol."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"263","DOI":"10.1016\/S0023-6438(02)00226-8","article-title":"Determination of in vitro antioxidant activity of fennel (Foeniculum vulgare) seed extracts","volume":"36","author":"Oktay","year":"2003","journal-title":"LWT Food Sci. Technol."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"3010","DOI":"10.1021\/jf0115589","article-title":"Thermal processing enhances the nutritional value of tomatoes by increasing total antioxidant activity","volume":"50","author":"Dewanto","year":"2002","journal-title":"J. Agric. Food Chem."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"654","DOI":"10.1016\/j.foodchem.2005.04.028","article-title":"Antioxidant activity of some Algerian medicinal plants extracts containing phenolic compounds","volume":"97","author":"Djeridane","year":"2006","journal-title":"Food Chem."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"873","DOI":"10.1016\/j.biortech.2004.09.006","article-title":"Polyphenol contents and antioxidant activity of Maydis stigma extracts","volume":"96","year":"2005","journal-title":"Bioresour. Technol."}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/18\/2743\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,8]],"date-time":"2026-09-08T10:15:42Z","timestamp":1788862542000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/18\/2743"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,8]]},"references-count":102,"journal-issue":{"issue":"18","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["plants15182743"],"URL":"https:\/\/doi.org\/10.3390\/plants15182743","relation":{},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,8]]}}}