Due to an increase in the number of herbicide-resistant weeds and environmental concerns about the use of synthetic herbicides, a great effort is being made in designing alternative weed management strategies. The present study was carried out in order to examine the impact of winter savory (Satureja montana) aqueous extract in natural weed management. We evaluated the effect of two concentrations (0.1 and 0.2%) of S. montana aqueous extract on the activity of antioxidant enzymes and the lipid peroxidation process in Jimson weed (Datura stramonium) seedlings. Our results showed that S. montana aqueous extract induced lipid peroxidation in roots of Jimson weed seedlings 72 hours after the treatment

Anačkov Goran

Malenčić Đorđe

Manojlović Ana

Prvulović Dejan

Šućur Jovana

Botanica Serbica

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42 (2): (2018) 271-275UDC: [[582.929.4:615.451.1]:632.954:632.5]:582.926.2DOI: 10.5281/zenodo.1468370Short Communication Received: 27 July 2017  Revision accepted: 30 January 2018© 2018 Institute of Botany and Botanical Garden Jevremovac, Belgrade✳correspondence: jovana.sucur@polj.edu.rsInfluence of winter savory (Satureja montana L.) aqueous extract on antioxidant properties of Jimson weed (Datura stramonium L.) Jovana Šućur1✳, Dejan Prvulović1, Ana Manojlović1, Goran Anačkov2  and Đorđe Malenčić1 1 University of Novi Sad, Faculty of Agriculture, 21000 Novi Sad, Serbia2 University of Novi Sad, Faculty of Science, Department of Biology and Ecology, 21000 Novi Sad, SerbiaABSTRACT: Due to an increase in the number of herbicide-resistant weeds and environmental concerns about the use of synthetic herbicides, a great effort is being made in designing alternative weed management strategies. The present  study was carried out in order to examine the impact of winter savory (Satureja montana) aqueous extract in natural weed management. We evaluated the effect of two concentrations (0.1 and 0.2%) of S. montana aqueous extract on the activity of antioxidant enzymes and the lipid peroxidation process in Jimson weed (Datura stramonium) seedlings. Our results showed that S. montana aqueous extract induced lipid peroxidation in roots of Jimson weed seedlings 72 hours after the treatment. Keywords: allelopathy, Datura stramonium, Satureja montanaSatureja montana L. (Lamiaceae) (winter savory) is a well known aromatic plant, often used in traditional medicine (SILVA et al. 2009). The aqueous extract of S. montana contains active compounds like the phenolic components caffeic acid and gallic acid (ŠUĆUR et al. 2015). As one of the major groups of compounds, plant phenolics possess a broad spectrum of chemical and biological activities (Zeng et al. 2001). The application of allelopathic interactions and natural substances in agricultural practice can reduce the use of synthetic pesticides in the management of weeds. Allelopathic interactions are based on the production of secondary biomolecules by plants which exert positive or negative effects on other plants (Macías et al. 2004). One of the main effects of released secondary biomolecules on tar-get plants is the excessive production of reactive oxygen species (ROS), molecules very toxic to cells (Bogatek & Gniazdowska 2007). Plants are able to cope with oxidative damage by using some antioxidant enzymes and free radical scavengers. Superoxide dismutase is the first enzyme in the detoxifying process; it catalyses the dismutation of superoxide free radical anions to mo-lecular oxygen and water, preventing the formation of other more toxic oxygen species such as the OH radical. Catalase and peroxidases metabolise hydrogen peroxide. The typical catalase reaction is the dismutation of two molecules of hydrogen peroxide to water and molecular oxygen (Mhamdi et al. 2010). Peroxidases catalyse the oxidation of a wide variety of substrates, using H2O2 or other peroxides (Hamid & Rehman 2009). Physiolog-ical activities are hindered by allelopathy (Siddique & Ismail 2013), and activity of antioxidant enzymes can thus be used as an indicator of oxidative stress in plants (Fakoorziba et al. 2014). One of the most resistant weed species is Jimson weed (Datura stramonium L.). Jim-son weed is a widespread annual plant, encountered in maize and sunflower fields. All parts of the plant are tox-ic, and the toxins in it are tropane belladonna alkaloids (Binev et al. 2006; Maheshwari et al. 2013). It inhibits the growth of surrounding plants through the mecha-272 vol. 42 (2)t 24 h 72 h 120 hLeavesCatalase Control 72.04 ± 6.75a      80.45 ± 1.33a       74.92 ± 5.55a        0.1 % 91.43 ± 14.66a        73.26 ± 13.01a       65.66 ± 2.08a       0.2 % 90.52 ± 5.30a        71.86 ± 10.27a       38.57 ± 4.69b       Superoxide dismutase Control 27.07 ± 1.02a      32.51 ± 0.15b        31.96 ± 0.43b       0.1 % 35.36 ± 0.92e      20.70 ± 0.17d        29.33 ± 0.21c       0.2 % 25.84 ± 0.23a     25.97 ± 0.09a            38.20 ± 0.56f       Guaiacol peroxidase Control (0.76 ± 0.03).103 a       (0.80 ± 0.09).103 a         (1.42 ± 0.09).103 c,d       0.1 % (0.82 ± 0.01).103 a      (1.12 ± 0.07).103 b         (1.25 ± 0.06).103 c        0.2 % (0.97 ± 0.05).103 a,b      (1.50 ± 0.11).103 d        (1.65 ± 0.04).103 d        Pyrogallol peroxidase Control (0.64 ± 0.06).103 a       (0.91 ± 0.077.86).103 a,c        (0.67 ± 0.05).103 a        0.1 % (0.79 ± 0.05).103 a,b,c       (1.02 ± 0.04).103 c       (1.32 ± 0.09).103 d        0.2 % (0.69 ± 0.08).103 a,b      (1.41 ± 0.13).103 d       (1.04 ± 0.03).103 c        MDA content Control 6.90 ± 1.57a,b 11.24 ± 2.08a,b 6.33 ± 0.09b0.1 % 9.88 ± 1.13a,b 9.07 ± 3.00a,b 6.32 ± 0.38b0.2 % 12.15 ± 2.44a 7.17 ± 0.61a,b 7.09 ± 0.14a,bRootsCatalase Control 23.64 ± 2.58a,b       21.93 ± 1.81a       17.86 ± 2.83a      0.1 % 34.43 ± 2.22a,b        32.89 ± 3.46a,b        57.38 ± 10.72b       0.2 % 29.29 ± 3.71a,b        46.77 ± 8.63a,b        80.27 ± 27.58c        Superoxide dismutase Control 68.79 ± 3.22a      57.16 ± 4.09a,b         96.37 ± 0.83c       0.1 % 103.57 ± 20.23c       63.75 ± 7.06a,b        99.96 ± 5.42c        0.2 % 39.77 ± 8.00b       54.06 ± 10.58a,b     118.97± 3.66c       Guaiacol peroxidase Control (4.58 ± 0.25).103 a,b       (5.60 ± 0.27).103 b,c        (6.76 ± 0.27).103 e        0.1 % (3.87 ± 0.46).103 a      (4.85 ± 0.10).103 b        (10.00 ± 0.22).103 f       0.2 % (2.93 ± 0.13).103 d       (5.80 ± 0.26).103 c       (3.92 ± 0.47).103 a       Pyrogallol peroxidase Control (4.36 ± 0.34).103 a      (5.61 ± 0.28).103 c        (3.77 ± 0.24).103 a,b        0.1 % (4.42 ± 0.19).103 a       (3.76 ± 0.64).103 a,b       (3.23 ± 0.09).103 b        0.2 % (3.63± 0.16).103 a,b       (4.28 ± 0.21).103 a        (3.25 ± 0.26).103 b        MDA content Control 8.22 ± 1.09a,b 8.15 ± 0.03a,b 7.36 ± 0.27a0.1 % 5.98 ± 0.17a 21.08 ± 7.05c 12.13 ± 0.64a,b,c0.2 % 23.99 ± 4.83d 18.34 ± 3.89b 12.34 ± 0.11a,b,cThe data are mean values ± standard error. a-fvalues without the same superscripts within each column differ significantly (P<0.05).Table 1. Effects of two concentrations (0.1 and 0.2%) of S. montana aqueous extract on the activity of antioxidant enzymes (U/mg of protein) and on MDA content (nmol/mg of protein) in leaves and roots of D. stramonium seedlings compared to the control group.273J. Šućur et al.: Influence of winter savory on Jimson weednism of allelopathy (Elisante & Ndakidemi 2014). It is known that root exudates are one of the main sources of allelochemicals released into the soil (Gatti et al. 2010). In line with this, it is very important to remove them from crops. The aim of this study was to investigate the impact of winter savory (S. montana) aqueous extract in natural weed management. The effect of S. montana aqueous extract on the lipid peroxidation process and activity of the antioxidant enzymes superoxide dismu-tase (SOD), catalase (CAT), guaiacol peroxidase, and py-rogallol peroxidase in leaves and roots of Jimson weed (D. stramonium) seedlings is examined in the study. Plant material. The wild aromatic plant S. montana was collected around the city of Podgorica in Montenegro (42º32’23.21’’ N, 19º20’02.17’’ E) at an elevation of 123 m a.s.l. in June of 2012 (voucher No. 2-1544). Datura stra-monium seeds were collected in Futog (Krndelja; UTM: 34T DR 2 01) in June of 2012 (Voucher No. 2-1486). Voucher specimens of both species are deposited in the herbarium of the Department of Biology and Ecology (BUNS), Faculty of Natural Sciences, University of Novi Sad (Holmgren & Holmgren 2003). Preparation of aqueous extract. Air–dried plant mate-rial (above-ground part) from S. montana was ground into powder. The powder (10 g) was extracted in 100 mL of boiling distilled water (10% w/v).Seedling growth. The experiment was performed at the Laboratory of Biochemistry, Faculty of Agriculture, Novi Sad, and conducted under controlled conditions (28°C, 60% relative humidity, 18-h photoperiod, and light intensity of 10,000 lx). Datura stramonium seeds were surface-sterilised with 3% H2O2 (v/v) and washed with sterilised deionised water. These seeds were placed in plastic pots containing sterile sand and maintained under dark conditions. Thirty-day-old seedlings were transplanted in plastic pots containing 700 mL of Ho-agland’s solution [10% MgSO4 × 7H2O, 10% Ca(NO3)2 × 4 H2O, 10% KH2PO4, 10% KNO3, microelements, 7.5% Fe-EDTA] and 7 (0.1 %) or 14 (0.2 %) mL of S. montana aqueous extract, while pots of the control contained the same volume of nutrient solution without addition of S. montana aqueous extract. Seedlings were harvested for determining the investigated biochemical parameters 24, 72, and 120 h after the treatments with S. montana aqueous extract.Biochemical assays. For determination of oxidative stress parameters, 2 g of fresh plant material (leaves and roots of D. stramonium plants, separately) was crushed and homogenised in 10 mL of phosphate buffer (0.1 M, pH 7.0). Homogenates were centrifuged for 20 min at 10.000 x g and filtered. The supernatants were used for biochemical assays. Lipid peroxidation was measured at 532 nm using the thiobarbituric acid (TBA) test (Man-dal et al. 2008). The total amount of TBA–reactive sub-stances was given in nmol of malondialdehyde (MDA) equivalents per mg of proteins. Catalase (CAT) (EC 1.11.1.6) activity was determined according to Sathya & Bjorn (2010). The decomposition of H2O2 was followed as the decrease in absorbance at 240 nm. Activity of the enzyme was expressed in U mg-1 of proteins. Superox-ide dismutase (SOD) (EC 1.15.1.1) activity was assayed according to a slightly modified version of the method of Mandal et al. (2008) by measuring its ability to in-hibit photochemical reduction of nitro blue tetrazolium (NBT) chloride. One unit of SOD activity was defined as the amount of the enzyme required to inhibit reduction of NBT by 50%.  Peroxidase (POD) (EC 1.11.1.7) activity was measured using guaiacol (guaiacol peroxidase) and pyrogallol (pyrogallol peroxidase) as substrates accord-ing to Morkunas & Gmerek (2007). Activity of the en-zyme was expressed in U mg-1 of proteins.Statistical analyses. All measurements were performed in triplicates. Values of the biochemical parameters were expressed as means ± standard error of the mean and tested by ANOVA, followed by comparison of the means by Duncan’s multiple range test (P<0.05). Data were ana-lysed using STATISTICA for Windows version 11.0. Allelopathic interactions and natural substances present in plant extracts are good candidates to be developed as bioherbicides. In the present study, a significant decrease of catalase activity was detected in the leaves of Jimson weed seedlings treated with a 0.2% concentration of S. montana aqueous extract 120 h after the treatment. A significant decrease of superoxide dismutase activity and significant increase of guaiacol and pyrogallol peroxi-dase activity were observed in the variants of treatments with both tested concentrations 72 h after the treatment (Table 1). In the roots of Jimson weed seedlings, a signifi-cant increase of catalase activity was recorded 120 h after treatment with 0.2% S. montana aqueous extract. Statis-tically significant increases in MDA accumulation were recorded in roots of Jimson weed 24 h after treatment with a 0.2% concentration and 72 h after treatment with a 0.1% concentration of S. montana aqueous extract. The results showed that the tested doses did not induce li-pid peroxidation in the leaves of Jimson weed seedlings. On the other hand, both concentrations induced higher accumulation of MDA,  an end-product of the lipid per-oxidation process, in the roots of Jimson weed seedlings during the first 72 h (Table 1). These results show that the roots were more affected than the leaves, the rea-son for this probably lying in fact that the roots were directly exposed to S. montana aqueous extract. Other studies have also shown the allelopathic potential of S. montana. Angelini et al. (2003) found that S. montana essential oil blocked germination of three weed species, 274 vol. 42 (2)viz., lambsquarters (Chenopodium album L.), little hog-weed (Portulaca oleracea L.), and barnyard grass [Echin-ochloa crus-galli (L.) Beauv.]. Furthermore, the aqueous extracts of other Satureja species were shown to have inhibitory effects on the percentage of weed seed germi-nation (Gholami et al. 2011).Our results showed that roots of Jimson weed were more affected by S. montana aqueous extract than leaves. Both tested concentrations of S. montana aqueous extract in-duced lipid peroxidation in Jimson weed roots 24 h and 72 h after the treatment. Sensitivity of D. stramonium to S. montana aqueous extract indicates that natural sub-stances present in plant extracts are good candidates to be developed as bioherbicides. REFERENCESAngelini LG, Carpanese G, Cioni PL, Morelli I, Macchia M & Flamini G. 2003. Essential oils from Mediterranean Lamiaceae as weed germination inhib-itors. Journal of Agricultural and Food Chemistry 51: 6158–6164.Binev R, Valchev I & Nikolov J. 2006. Hematological studies on Jimson weed (Datura stramonium) intoxi-cation in horses. Trakia Journal of Sciences 4(1): 43–48.Bogatek R & Gniazdowska A. 2007. ROS and phyto-hormones in plant-plant allelopathic interaction. Plant Signaling and Behavior 2(4): 317–318.Elisante F & Ndakidemi PA. 2014. Allelopathic effect of Datura stramonium on the survival of grass and leg-ume species in the conservation areas. American Jour-nal of Research Communication 2(1): 27–43.Fakoorziba MR, Moemenbellah-Fard MDJ, Azizi K, Shekarpoor H & Alipoor H. 2014. Excito-repel-lency effects of Salvia sclarea L. (Lamiaceae) extracts on adult house flies, Musca domestica L. (Diptera: Muscidae). Journal of Health Sciences and Surveillance System 2(1): 2–7.Gatti AB, Ferreira AF, Ardun M & Perez CGA. 2010. Allelopathic effects of aqueous extracts of Arti-stolochia esperanzae O. Kuntze on developemment of Sesamum indicum L. seedlings. Acta Botanica Brasili-ca 24(2):454-461.Gholami BA, Faravani M & Kashki MT. 2011. Allelo-pathic effects of aqueous extract from Artemisia kopet-daghensis and Satureja hortensis on growth and seed germination of weeds. Journal of Applied Environmen-tal and Biological Sciences 1(9): 283–29.Hamid M & Rehman K. 2009. Potential applications of peroxidases. Food Chemistry 115: 1177–1186.Holmgren PK & Holmgren NH. 2003. Additions to Index Herbariorum (Herbaria), Edition – 8 Four-teenth Series. Taxon 52: 385-389. http://sciweb.nybg.org/science2/IndexHerbariorum.asp Macías FA, Galindo JCG, Molinillo JMG & Gutler HG. 2004. Allelopathy. Chemistry and mode of action of allelochemicals. CRC Press, NY.Maheshwari NO, Khan A & Chopade AB. 2013. Re-discovering the medicinal properties of Datura sp.: A review. Journal of Medicinal Plants Research 7(39): 2885–2897.Mandal S, Mitra A, Mallick N. 2008. Biochemical characterization of oxidative burst during interaction between Solanum lycopersicum and Fusarium ox-ysporum f. sp. lycopersici. Physiological and Molecular Plant Pathology 72: 56–61.Mhamdi A, Queval G, Chaouch S, Vanderauwera S, van Breusegem F & Noctor G. 2010. Catalase function in plants: a focus on Arabidopsis mutants as stress-mimic models. Journal of Experimental Botany 61: 4197-4220.Morkunas I & Gmerek J. 2007. The possible involve-ment of peroxidase in defense of yellow lupine embryo axes against Fusarium oxysporum. Journal of Plant Physiology 164: 185–194.Sathya E & Bjorn M. 2010. Plant stress tolerance. Hu-mana Press, Oklahoma.Siddique MAB & Ismail BS. 2013. Allelopathic effects of Fimbristylis miliacea on the physiological activities of five Malaysian rice varieties. Australian Journal of Crop Science 7(13): 2062–2067.Silva FVM, Martins A, Salta J, Neng NR, Nogue-ira JMF, Mira D, Gaspar N, Justino J, Grosso C, Urieta JS, Palavra AMS & Rauter AP. 2009. Phy-tochemical profile and anticholinesterase and antimi-crobial activities of supercritical versus conventional extracts of Satureja montana. Journal of Agricultural and Food Chemistry 57: 11557–11563.Šućur J, Popović A, Petrović M, Anačkov G, Bursić V, Kiprovski B & Prvulović D. 2015. Allelopathic ef-fects and insecticidal activity of the aqueous extract of Satureja montana L. Journal of the Serbian Chemical Society 80(4): 475–484.Zeng RS, Luo SM, Shi YH, Shi MB & Tu CY. 2001. Physiological and biochemical mechanism of allelop-athy of secalonic acid F on higher plants. Agronomy Journal 93(1): 72–79.275J. Šućur et al.: Influence of winter savory on Jimson weedZbog sve veće otpornost korovskih vrsta prema herbicidima, kao i problema koji se javljaju nakon upotrebe sintetičkih herbicida, ulažu se veliki napori u izradi alternativnih strategija upravljanja korovom. Ovo istraživanje je sprovedeno kako bi se ispitao uticaj vodenog ekstrakta rtanjskog čaja (Satureja montana) kao prirodnog herbicida u upravljanju korovom. Ispitan je uticaj dve koncentracije (0.1 i 0.2%) vodenog ekstrakta S. montana na aktivnost antioksidativnih enzima i proces lipidne peroksidacije u sadnicama tatule (Datura stra-monium). Dobijeni rezultati su pokazali da je povećanje intenziteta lipidne peroksidacije zabeleženo u korenu tatule 72 časa nakon tretmana. Ključne reči: alelopatija, Datura stramonium, Satureja montanaUticaj vodenog ekstrakta Satureja montana L. na an-tioksidativna svojstva tatule (Datura stramonium L.)Jovana Šućur, Dejan Prvulović, Ana Manojlović, Goran Anačkov  i Đorđe MalenčićREzImE276 vol. 42 (2)

Influence of winter savory (Satureja montana L.) aqueous extract on antioxidant properties of Jimson weed (Datura stramonium L.)

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Influence of winter savory (Satureja montana L.) aqueous extract on antioxidant properties of Jimson weed (Datura stramonium L.)

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