{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,12,24]],"date-time":"2025-12-24T14:47:34Z","timestamp":1766587654864,"version":"build-2065373602"},"reference-count":46,"publisher":"MDPI AG","issue":"24","license":[{"start":{"date-parts":[[2022,12,15]],"date-time":"2022-12-15T00:00:00Z","timestamp":1671062400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>The chemical profile of the essential oil (EO) of the aerial parts of Thymus leucotrichus var. creticus (Lamiaceae), a taxon not previously studied, was investigated by GC\u2013MS analysis, using a DB\u2013Wax polar column. Oxygenated monoterpenes and monoterpene hydrocarbons dominate the EO, with thymol (46.97%) and p-cymene (28.64%) as the main constituent of these two classes, respectively. The ability of the EO of T. leucotrichus to reduce Cd toxicity was studied in aquatic moss Leptodictyum riparium. To study EO-induced tolerance to Cd toxicity, apex growth, number of dead cells, DNA damage and antioxidant response in gametophytes were examined. The exogenous application of the EO yields a resumption of growth rate and a reduction in the number of dead cells; it also reduces the oxidative stress induced by Cd, as demonstrated by the reduction of the ROS content (with a decrease of 1.52% and 5%) and by the increased activity of antioxidant enzymes such as superoxide dismutase (SOD) (with an increase of 1.44% and 2.29%), CAT catalase (1.46% and 2.91%) and glutathione-S-transferase GST (1.57% and 1.90%). Furthermore, the application of the EO yields a reduction of DNA damage. These results clearly indicate the protective capacity of the EO of T. leucotrichus in modulating the redox state through the antioxidant pathway by reducing the oxidative stress induced by Cd.<\/jats:p>","DOI":"10.3390\/plants11243529","type":"journal-article","created":{"date-parts":[[2022,12,15]],"date-time":"2022-12-15T02:20:57Z","timestamp":1671070857000},"page":"3529","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":8,"title":["Chemical Composition of Thymus leucotrichus var. creticus Essential Oil and Its Protective Effects on Both Damage and Oxidative Stress in Leptodictyum riparium Hedw. Induced by Cadmium"],"prefix":"10.3390","volume":"11","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-4656-320X","authenticated-orcid":false,"given":"Viviana","family":"Maresca","sequence":"first","affiliation":[{"name":"Department of Biology, University of Naples Federico II, 80126 Naples, Italy"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7961-3894","authenticated-orcid":false,"given":"Natale","family":"Badalamenti","sequence":"additional","affiliation":[{"name":"Department of Biological, Chemical and Pharmaceutical Sciences and Technologies (STEBICEF), Universit\u00e0 degli Studi di Palermo, Viale delle Scienze, Ed. 17, 90128 Palermo, Italy"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Vincenzo","family":"Ilardi","sequence":"additional","affiliation":[{"name":"Department of Biological, Chemical and Pharmaceutical Sciences and Technologies (STEBICEF), Universit\u00e0 degli Studi di Palermo, Viale delle Scienze, Ed. 17, 90128 Palermo, Italy"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0583-0487","authenticated-orcid":false,"given":"Maurizio","family":"Bruno","sequence":"additional","affiliation":[{"name":"Department of Biological, Chemical and Pharmaceutical Sciences and Technologies (STEBICEF), Universit\u00e0 degli Studi di Palermo, Viale delle Scienze, Ed. 17, 90128 Palermo, Italy"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Paola","family":"Bontempo","sequence":"additional","affiliation":[{"name":"Department of Precision Medicine, University of Campania \u201cLuigi Vanvitelli\u201d, Via L. De Crecchio 7, 80138 Naples, Italy"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Adriana","family":"Basile","sequence":"additional","affiliation":[{"name":"Department of Biology, University of Naples Federico II, 80126 Naples, Italy"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1968","published-online":{"date-parts":[[2022,12,15]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"164","DOI":"10.1016\/j.ecoenv.2014.03.007","article-title":"Citric acid assisted phytoremediation of cadmium by Brassica napus L.","volume":"106","author":"Ehsan","year":"2014","journal-title":"Ecotox. Environ. Saf."},{"key":"ref_2","first-page":"872875","article-title":"Molecular mechanism of heavy metal toxicity and tolerance in plants: Central role of glutathione in detoxification of reactive oxygen species and methylglyoxal and in heavy metal chelation","volume":"2012","author":"Hossain","year":"2012","journal-title":"J. Bot."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"421","DOI":"10.1016\/S1002-0160(17)60339-4","article-title":"Morphological and physiological responses of plants to cadmium toxicity: A review","volume":"27","author":"Shanying","year":"2017","journal-title":"Pedosphere"},{"key":"ref_4","unstructured":"He, S., He, Z., Yang, X., Stoffella, P.J., and Baligar, V.C. (2015). Advances in Agronomy, Elsevier."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Jiang, Y., Fan, M., Hu, R., Zhao, J., and Wu, Y. (2018). Mosses are better than leaves of vascular plants in monitoring atmospheric heavy metal pollution in urban areas. Int. J. Environ. Res. Public Health, 15.","DOI":"10.3390\/ijerph15061105"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"401","DOI":"10.1016\/j.envpol.2007.07.004","article-title":"Comparison of the heavy metal bioaccumulation capacity of an epiphytic moss and an epiphytic lichen","volume":"151","author":"Basile","year":"2008","journal-title":"Environ. Pollut."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Bellini, E., Maresca, V., Betti, C., Castiglione, M.R., Fontanini, D., Capocchi, A., Sorce, C., Bors\u00f2, M., Bruno, L., and Sorbo, S. (2020). The moss Leptodictyum riparium counteracts severe cadmium stress by activation of glutathione transferase and phytochelatin synthase, but slightly by phytochelatins. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21051583"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"665","DOI":"10.1016\/j.ecoenv.2018.07.122","article-title":"Functional and structural biomarkers to monitor heavy metal pollution of one of the most contaminated freshwater sites in Southern Europe","volume":"163","author":"Maresca","year":"2018","journal-title":"Ecotox. Environ. Saf."},{"key":"ref_9","unstructured":"(2022, September 22). Plants of the World Online. Available online: https:\/\/powo.science.kew.org\/."},{"key":"ref_10","first-page":"144","article-title":"Flavonoids and further polyphenols in the genus Thymus","volume":"Volume 17","year":"2002","journal-title":"Thyme: The Genus Thymus"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"287","DOI":"10.1016\/j.tifs.2019.01.020","article-title":"Thymus spp. plants\u2013Food applications and phytopharmacy properties","volume":"85","author":"Salehi","year":"2019","journal-title":"Trends Food Sci. Technol."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"16","DOI":"10.1016\/j.jep.2009.06.021","article-title":"Traditional uses of plants in the Eastern Riviera (Liguria, Italy)","volume":"125","author":"Cornara","year":"2009","journal-title":"J. Ethnopharmacol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"e1900254","DOI":"10.1002\/cbdv.201900254","article-title":"Review Traditional Uses, Chemical constituents and biological activities of plants from the Genus Thymus","volume":"16","author":"Li","year":"2019","journal-title":"Chem. Biodiv."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"307","DOI":"10.1016\/0003-9861(78)90039-5","article-title":"Biosynthesis of aromatic monoterpenes: Conversion of \u03b3-terpinene to p-cymene and thymol in Thymus vulgaris L.","volume":"187","author":"Poulose","year":"1978","journal-title":"Arch. Biochem. Biophys."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"400","DOI":"10.1016\/0003-9861(78)90104-2","article-title":"\u03b3-Terpinene synthetase: Akey enzyme in the biosynthesis of aromatic monoterpenes","volume":"191","author":"Poulose","year":"1978","journal-title":"Arch. Biochem. Biophys."},{"key":"ref_16","first-page":"75","article-title":"Essential oil chemistry of the genus Thymus\u2014A global view","volume":"Volume 17","year":"2002","journal-title":"Thyme: The Genus Thymus"},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Peter, K.V. (2004). Thyme in Handbook of Herbs and Spices, Woodhead Publish.","DOI":"10.1533\/9781855738355"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1016\/j.materresbull.2018.09.025","article-title":"Biocompatibility assessment and photocatalytic activity of bio-hydrothermal synthesis of ZnO nanoparticles by Thymus vulgaris leaf extract","volume":"109","author":"Zare","year":"2019","journal-title":"Mater. Res. Bull."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"508","DOI":"10.1016\/j.ijbiomac.2018.10.042","article-title":"Cellulose acetate\/AgNPs-organoclay and\/or thymol nano-biocomposite films with combined antimicrobial\/antioxidant properties for active food packaging use","volume":"121","author":"Dairi","year":"2019","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"1379","DOI":"10.2217\/fmb-2019-0291","article-title":"Morpho-mechanical and antimicrobial properties of Coridothymus capitatus L. essential oil loaded into ultrafine poly (lactic acid) fibers prepared by electrospinning","volume":"15","author":"Scaffaro","year":"2020","journal-title":"Future Microbiol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"444","DOI":"10.1016\/j.jcis.2016.11.048","article-title":"Evaluation of Thymus vulgaris plant extract as an eco-friendly corrosion inhibitor for stainless steel 304 in acidic solution by means of electrochemical impedance spectroscopy, electrochemical noise analysis and density functional theory","volume":"490","author":"Ehsani","year":"2017","journal-title":"J. Colloid Interface Sci."},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"D\u2019Agostino, G., Badalamenti, N., Giambra, B., Palla, F., and Bruno, M. (2021). The application of the essential oils of Thymus vulgaris L. and Crithmum maritimum L. as biocidal on two Tholu Bommalu Indian leather puppets. Plants, 10.","DOI":"10.3390\/plants10081508"},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Tutin, T.G., Heywood, V.H., Burges, N.A., Moore, D.M., Valentine, D.H., Walters, S.M., and Webb, D.A. (1973). Flora Europaea, Cambridge University Press.","DOI":"10.2307\/1218149"},{"key":"ref_24","unstructured":"(2022, October 10). Euro + Med. Available online: https:\/\/ww2.bgbm.org\/EuroPlusMed\/query.asp."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"3849","DOI":"10.1021\/jf021232f","article-title":"Chemical composition and antibacterial activity of the essential oils of Thymus spinulosus Ten. (Lamiaceae)","volume":"51","author":"Bruno","year":"2003","journal-title":"J. Agric. Food Chem."},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Badalamenti, N., Vaglica, A., Maggio, A., and Bruno, M. (2022). A new ferulol derivative isolated from the aerial parts of Ferulago nodosa (L.) Boiss. growing in Sicily (Italy). Nat. Prod. Res., in press.","DOI":"10.1080\/14786419.2022.2074995"},{"key":"ref_27","doi-asserted-by":"crossref","unstructured":"Di Napoli, M., Maresca, V., Varcamonti, M., Bruno, M., Badalamenti, N., Basile, A., and Zanfardino, A. (2020). (+)-(E)-Chrysanthenyl acetate: A molecule with interesting biological properties contained in the Anthemis secundiramea (Asteraceae) flowers. Appl. Sci., 10.","DOI":"10.3390\/app10196808"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"243","DOI":"10.3892\/mmr.2022.12759","article-title":"Essential oil of Foeniculum vulgare subsp. piperitum fruits exerts an anti\u2014tumor effect in triple\u2014negative breast cancer cells","volume":"26","author":"Lauricella","year":"2022","journal-title":"Mol. Med. Rep."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"1097","DOI":"10.1080\/0972060X.2021.2003256","article-title":"Comparison of volatile constituents, antioxidant and antimicrobial activities of Thymus leucotrichus (Lamiaceae) stem and leaves essential oils from both natural resources and in vitro derived shoots","volume":"24","year":"2021","journal-title":"J. Essent. Oil. Bear. Plants"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"e2100498","DOI":"10.1002\/cbdv.202100498","article-title":"Essential oil composition of five Thymus species from Bulgaria","volume":"18","author":"Trendafilova","year":"2021","journal-title":"Chem. Biodiv."},{"key":"ref_31","first-page":"419","article-title":"Correlation between chemical composition of Greek essential oils and their antibacterial activity against food-borne pathogens","volume":"2","author":"Chorianopoulos","year":"2007","journal-title":"Nat. Prod. Commun."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1002\/lipi.19860880210","article-title":"Biochemical studies on the essential oils of some medicinal plants","volume":"88","author":"Farag","year":"1986","journal-title":"Fette. Seifen. Anstrichmitt."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1002\/ffj.2730040311","article-title":"Essential oil of Thymus glandulosus Lag. ex H. del Villar","volume":"4","author":"Adzet","year":"1989","journal-title":"Flav. Frag. J"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"4614","DOI":"10.3390\/molecules14114614","article-title":"Chemical composition and antimicrobial activity of the essential oils from two species of Thymus growing wild in Southern Italy","volume":"14","author":"Bruno","year":"2009","journal-title":"Molecules"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"6314","DOI":"10.1021\/jf0606104","article-title":"Composition of the essential oils of Thymus and Origanum species from Algeria and their antioxidant and antimicrobial activities","volume":"54","author":"Hazzit","year":"2006","journal-title":"J. Agric. Food. Chem."},{"key":"ref_36","first-page":"1251","article-title":"Thymus ciliatus\u2014The highest thymol containing essential oil of the genus","volume":"4","author":"Kabouche","year":"2009","journal-title":"Nat. Prod. Commun."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"577","DOI":"10.1080\/0972060X.2014.901616","article-title":"Chemical composition and antimicrobial activity of the volatile oils from leaf, flower, stem and root of Thymus transcaucasicus from Iran","volume":"17","author":"Ezzatzadeh","year":"2014","journal-title":"J. Essent. Oil Bear. Plants"},{"key":"ref_38","first-page":"80","article-title":"study on chemical composition and antimicrobial activity of the essential oil of Thymus trautvetteri Klokov & Desj.\u2013Shost","volume":"3","author":"Shahnazi","year":"2007","journal-title":"J. Med. Plants"},{"key":"ref_39","first-page":"34","article-title":"Composition of the essential oils of Thymus deanensis Celak var. deanensis from different regions of Iran","volume":"1","author":"Abousaber","year":"2012","journal-title":"J. Med. Plants"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"385","DOI":"10.1080\/0972060X.2010.10643839","article-title":"Chemical composition of the essential oil of Thymus migricus Klokov & Desj. -Shost. from Iran","volume":"13","author":"Yavari","year":"2010","journal-title":"J. Essent. Oil Bear. Plants"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"169","DOI":"10.1016\/j.mrrev.2008.07.003","article-title":"Repair and tolerance of oxidative DNA damage in plants","volume":"681","author":"Ariza","year":"2009","journal-title":"Mutat. Res."},{"key":"ref_42","unstructured":"European Pharmacopoeia (2008). Determination of essential oils in herbal drugs. Eur. Pharm., 307, 251\u2013256."},{"key":"ref_43","first-page":"1","article-title":"GC and GC\u2014MS Analysis of volatile compounds from Ballota nigra subsp. uncinata collected in Aeolian Islands, Sicily (Southern Italy)","volume":"15","author":"Rigano","year":"2020","journal-title":"Nat. Prod. Commun."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1007\/s00709-003-0028-0","article-title":"Accumulation, localisation, and toxic effects of cadmium in the liverwort Lunularia cruciata","volume":"223","author":"Carginale","year":"2004","journal-title":"Protoplasma"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"374","DOI":"10.1080\/15226514.2011.620653","article-title":"Toxicity, accumulation, and removal of heavy metals by three aquatic macrophytes","volume":"14","author":"Basile","year":"2012","journal-title":"Int. J. Phytoremed."},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Esposito, S., Loppi, S., Monaci, F., Paoli, L., Vannini, A., Sorbo, S., Maresca, V., Fusaro, L., Karam, E.A., and Lentini, M. (2018). In-field and in-vitro study of the moss Leptodictyum riparium as bioindicator of toxic metal pollution in the aquatic environment: Ultrastructural damage, oxidative stress and HSP70 induction. PLoS ONE, 13.","DOI":"10.1371\/journal.pone.0195717"}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/11\/24\/3529\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T01:41:40Z","timestamp":1760146900000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/11\/24\/3529"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,12,15]]},"references-count":46,"journal-issue":{"issue":"24","published-online":{"date-parts":[[2022,12]]}},"alternative-id":["plants11243529"],"URL":"https:\/\/doi.org\/10.3390\/plants11243529","relation":{},"ISSN":["2223-7747"],"issn-type":[{"type":"electronic","value":"2223-7747"}],"subject":[],"published":{"date-parts":[[2022,12,15]]}}}