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Thymus contra Fusarium oxysporum Boualem Boumaaza 1,2 * M’hamed Benada 3 Sofiane Boudalia 4 Ibrahim E. Benzohra 5 Abdelhamid Gacemi 6 Omar Khaladi 4 Mohamed Benkhelifa 1 Rev. Fac. Agron. (LUZ). 2022, 39(3): e223941 ISSN 2477-9407 DOI: https://doi.org/10.47280/RevFacAgron(LUZ).v39.n3.07 Crop Production Associate editor: Dra. Lilia Urdaneta University of Zulia, Faculty of Agronomy Bolivarian Republic of Venezuela 1 Department of Agronomy, University of Abdelhamid Ibn Badis, BP 300, 27000, Mostaganem, Algeria. 2 University of Ibn khaldounTiaret. BP 78 zaâroura 14000, Tiaret, Algéria. 3 University of Ahmed Zabana. BP 48000 Bormadia Relizane. 4 Laboratory of Biology, Water and Environment (LBEE), University of 8th May, 1945 Guelma, BP 401 24000 Guelma, Algeria. 5 Scientific and Technical Research Center on Arid Regions (CRSTRA) BP. Box 1682 RP 07000, Biskra, Algeria. Conservation Laboratory Wetlands. 6 INRA Station de Recherche Hmadna. BP 48017 Relizane. Received: 01-06-2022 Accepted: 02-08-2022 Published: 22-082022 Abstract Thyme ( Thymus sp.), a medicinal plant of the family Lamiaceae , is used in traditional medicine, contains a wide array of medicinally active components, in their great majority of a rather complex mixture of thymol, ρ-Cymene, γ-Terpinene, β-Caryophyllen, etc. This study aimed to evaluate the efficacy of Thymus vulgaris extract against Fusarium oxysporum f. sp. radicis-lycopersici strain under saline conditions, assuming soil with high salt content of the arid regions. Essential oil was extracted by hydrodistillation technique using a Clevenger apparatus. The essential oil compounds were identified by GC-MS analysis. Antifungal activity of essential oil against Fusarium oxysporum f. sp . radicis-lycopersici was investigated by agar dilution method. The main constituents of thyme essential oil were thymol (76.96 %), ρ-cymene (9.89 %) and γ-Terpinene (1.92 %). Essential oil from Thymus resented high in vitro activity, in controlling conidial germination and mycelial growth. However, the oil was significantly not active against the spore production under a salinity medium. The results showed that mycelial growth was stimulated in concentrations with 0.6-1.5 %. In contrast, it was significantly reduced at a higher concentration (2 %). The application of NaCl caused a significant increase in the conidia production at various concentrations tested. NaCl has a minor inhibitory effect on conidial germination only when the concentration was 2 %. 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Rev. Fac. Agron. (LUZ). 2022, 39(3): e223941. July - September. ISSN 2477-9407. 2-6 | Resumen El tomillo ( Thymus sp.), una planta de la familia Lamiaceae , se utiliza en la medicina tradicional, contiene una amplia gama de componentes medicinalmente activos, en su gran mayoría de una mezcla bastante compleja de timol, ρ-cimeno, γ-terpineno y β-Cariofileno, entre otros. El objetivo del presente estudio fue evaluar la eficacia del extracto de Thymus vulgaris contra Fusarium oxysporum f. sp. radicis-lycopersici bajo condiciones salinas, simulando suelos con altos contenidos de sales de las regiones áridas. El aceite esencial se extrajo mediante hidrodestilación utilizando el aparato Clevenger. Los compuestos del aceite esencial se identificaron mediante el análisis GC-MS. La actividad antifúngica del aceite se evaluó mediante el método de dilución en agar. Los principales componentes del aceite esencial del tomillo fueron timol (76,96 %), ρ-cimeno (9,89 %) y γ-terpineno (1,92 %). El aceite presentó alta actividad in vitro , en el control de la germinación de los conidios y del crecimiento micelial. Sin embargo, el aceite no inhibió la esporulación en el medio salino. El crecimiento micelial fue estimulado al 0,6-1,5 %, pero se redujo significativamente a la concentración más alta (2 %). La aplicación de NaCl provocó un aumento significativo en la producción de conidios en todas las concentraciones. La germinación conidial fue inhibida ligeramente por el NaCl solo al 2 %. Los resultados indican que la salinidad disminuye el potencial antifúngico del aceite esencial de Thymus contra el patógeno. Palabras clave: Actividad antimicrobiana, biocontrol, extractos de plantas, planta medicinal, cloruro de sodio. Resumo O tomilho ( Thymus sp.), planta medicinal da família Lamiaceae , utilizada na medicina tradicional, contém uma grande variedade de componentes medicinais ativos, em sua grande maioria uma mistura bastante complexa de Timol, ρ-Cymene, γ-Terpinene, β-Caryophyllen, etc. O presente estudo foi avaliar a eficácia do extrato de Thymus vulgaris contra Fusarium oxysporum f. sp . radicis-lycopersici sob condições salinas, assumindo solos com alto teor de sal das regiões áridas. O óleo essencial por extraído pe la técnica de hidrodestilação utilizando aparelho de Clevenger. O composto de óleo essencial por identificado por análise GC-MS. Atividade antifúngica do óleo esencial contra Fusarium oxysporum f. sp . radicis-lycopersici por investigado pelo método de diluição em agar. O principais constituintes do óleo esencial de tomilho foram timol (76.96 %), ρ-cimeno (9.89 %) e γ-Terpineno (1.92 %). O óleo esencial de Timo apresentou alta ativida de in vitro , no control da germinação de conídios e do crescimento micelial. No entanto, o óleo não por significativamente ativo contra produção de esporos meio de salinidade. Os resultados sugerem que o crescimento micelial por estimulado em concentrações de 0,6-1.5 %. Pelo contrário, reduziu significativamente na concentração mais alta (2 %). A aplicação de NaCl causou um aumento significativo na produção de conidiogênese em várias concentrações testadas. A germinação dos conídios for levemente inibida pelo cloreto de sódio somente quando a concentração for de 2 %. Os resultados este estudo indicaque a salinidade diminui a eficácia do óleo esencialcontra o patógeno. Palavras-chave: Actividad antimicrobiana, biocontrole, extratos de plantas, planta medicinal, cloreto de sódio. Introduction Fusarium oxysporum is an ubiquitous fungus, widely distributed among soil, plants, plant debris and other organic substrates. This fungus is one of the most important phytopathogenic and toxigenic fungi, affecting the health and survivability of plants in more than 100 different crops (Al-Hatmi et al ., 2016). Additionally, Fusarium oxysporum has numerous specialized forms (f. sp.) that infect a range of host plants causing diseases such as vascular wilt, corm rot, and root rot (Edel-Hermann and Lecomte, 2019). Another important characteristic of the species of this genus is their ability to produce a various of mycotoxins that can play an important role in the pathogenesis (Lombard et al ., 2019). Preventive measures, such as chemical pesticides have controlled the disease to some extent. However, excessive application of fungicides has led to severe damage to soil microbial communities and fertility and may leave behind toxic residues in treated products (Meena et al ., 2020). Additionally, intensive use of fungicides can result in the development of pathogens acquired resistance to the fungicide (Corkley et al ., 2021). Due to the problems caused by chemical substances, the development of alternative control measures is of great importance. Biological control has been considered as a desirable and realistic alternative. Numerous studies have demonstrated the ability of several essential oils to possess antibacterial, antifungal, antiviral, antioxidant activities and play an important role in the protection of the plants against plant pathogens both in vitro and in vivo (Mutlu- Ingok et al ., 2020; Marín-Tinoco et al ., 2021). Note that essential oils can be an effective solution, as their toxicity is much lower, better specificity of action, biodegradable and environmentally friendly (Campos et al ., 2019). Being volatile, they can act as fumigants, repellents, and contact insecticides or as reproduction inhibitors (Gao et al ., 2020; Silva-Marrufo and Marín-Tinoco, 2021). Essential oils interact with the odor receptors of insect pests ( Ceratitis capitata ), triggering various behaviors: flight, attraction, oviposition, etc. (Benelli et al ., 2012). The studies published on the activity of essential oils as herbicides are numerous and generally cover seed germination inhibition tests (Yilar et al., 2020). Those that appear to be the most active are essential oils containing phenols (thymol, carvacrol), ketones (carvone, pulegone) or etheroxides (eucalyptol or 1,8-cineol). Plants of the genus Thymus are widely used in food flavouring, as well as aroma additives, in perfumery, in folk medicine and in pharmacological sector as natural antioxidants and antimicrobial agents (Gema, 2020). In North Africa, most of the thyme produced is in the form of dried herbs. With mean annual rainfalls of 150-400 mm.years -1 , the yield of Thymus vulgaris as fresh herbs may be 5 to 6 t.ha -1 with 2 t.ha -1 for the production of dried herbs. With an irrigated system, in the presence of fertilization, the yield can be 9.77 t.ha -1 , with an oil recovery yield of 43.14 to 48.9 kg.ha -1 (Kozera et al ., 2015). It has been reported that essential oils possess various actions on microorganisms, such as disruption of the cytoplasmic membrane, disruption of the proton driving force, electron leakage and coagulation of the protein content of cells, acidification from inside the cell, blocking the production of cellular energy and the synthesis of structural components. Thymol and carvacrol have antimicrobial activity, which is related to inhibition of membrane permeability, resulting in ions and ATP leakage, inhibition ergosterol biosynthesis and cell death (Alizadeh et al ., 2018). This scientific publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818. 3-6 | Antimicrobial activity of essential oils is strongly affected by many factors such as high temperature, poor water solubility, light, and oxygen (de Souza et al ., 2008; Turek and Stintzing, 2012). This study aimed to assess the antifungal activity of the essential oil from Thymus against Fusarium oxysporum f. sp. radicis-lycopersici in growth media amended with various levels of sodium chloride. Materials and methods Inoculum preparation The tested Fusarium oxysporum f. sp. radicis-lycopersici strain M27b27245 was preserved at the Plant Protection Laboratory of the Agronomy Science Department, University of Abdelhamid Ibn Badis, Mostaganem, Algeria. Fusarium oxysporum readily produced conidia after 14 days on potato dextrose agar (PDA; potato dextrose agar. SPA CRAPC Algeria) plate at 25 ° C in the dark. Conidial suspension was obtained by flooding plates with distilled water and rubbing gently with a glass rod, then filtering through sterile cheesecloth. The conidia concentration present in the initial suspension (adjusted to the concentration 1x10 5 conidia.mL -1 ) was quantified using the hemocytometer. Analysis of the essential oil of Thymus vulgaris Thymus vulgaris was supplied for the study by the National Agronomic Institute of Tunisia (INAT), and the experiment was conducted from December 2017 to January 2018. Hydro- distillationwascarried out using a Clevenger-type device according to the method recommended in British Pharmacopoeia (British Pharmacopoeia, 2016), where 500 g of vegetable matter is introduced with 3 L water into a 5 L flask. After installation and closing of the assembly, the flask heater is started up with an optimum adjustment of the heating to allow stability of the extraction at a constant and well- controlled speed. The vapor charged with essential oil arrives in the condenser. The total duration of the extraction was estimated at 3 h. The essential oil differs from the hydrosol by its difference in density and color. It is separated from the by decantation. It is then dried with anhydrous sodium sulfate (Na 2 SO 4 ) then recovered and stored in a cool place (4 °C). Oil extracted was analyzed by GC/MS using a gas chromatograph (HP 5890-Serie II) coupled to a mass spectrometer (HP-MSD 5972 A) with an HP-5MS capillary column (30 m x 0. 25 mm, film thickness, 0.25 µm). The temperature of the column was programmed at 50 ° C for 1 min, then 7 ° C / min at 250 ° C, and finally left at 250 ° C for 5 min. The helium injection (1 mL.min -1 ). The temperature of the injector port was 240 ° C, while that of the detector set at 250 ° C. The ionizing energy was 70 eV, the full-scan mass spectra: from 40 to 500 amu. The identification of the components was carried out on the basis of chromatographic retention indices and by comparison of the mass spectra recorded with the calculated spectral library (Al-Asmari et al ., 2017). In vitro antifungal assay Effects on mycelial growth The antifungal activity of the essential oil against to Fusarium oxysporum f. sp. radicis-lycopersici was investigated by agar dilution method (Souza et al ., 2002). In sterile Petri dishes with 9 cm diameter containing 20 mL potato dextrose agar (PDA) mixed with five levels of NaCl to obtain final concentrations 0.3, 0.6, 1, 1.5 and 2 %. Then, 100 μL of the essential oil was evenly spread on the surface of Petri dishes containing PDA + NaCl. Essential oil was previously diluted, to then adjust the concentration to 1 %. 30 μL of the fungi Boumaaza et al. Rev. Fac. Agron. (LUZ). 2022, 39(3): e223941 suspension 105 conidia.mL - 1 was placed at the center of each Petri plate. In this assay, an amount of essential oil was added at minimum inhibitory concentration (MIC) (1 %). Cultures were incubated at 25 °C in the dark. Colony diameters were measured in two perpendicular directions, after 24 h and again after 96 h. Each treatment had five replications. In the control group, sterile distilled water was used in place of the oil, and no additional NaCl was added. Effects on sporulation C o n i d i a s u s p e n s i o n o f Fusarium oxysporum f. sp. radicis- lycopersici was obtained from two weeks-old culture incubated at 25 ° C in the dark. Plates were flooded with sterile-distilled water using 0.5 % (v/v) Tween 80 solution and conidia were gently dislodged from the mycelium using a sterile magnetic stir bar placed on the agar and set stirring for 5 min to loosen the conidia. The suspension was filtered through a glass filter to eliminate mycelium. Finally, the conidia concentration was quantified using the hemocytometer. Effects on the germination of conidia To determine the influence of NaCl upon the antifungal activity of the essential oil on spore germination of Fusarium oxysporum f. sp. radicis-lycopersici , a drop containing 100 conidia was transferred on to water agar plates enriched with NaCl and essential oil. The plates were incubated at 25 ° C in the dark for 24 h. The results were expressed as the percentage of germinated conidia observed under the optical microscope at 400x magnification. The conidium was considered as germinated if the germ tube length was at least twice the length of the conidium. All results are presented as mean ± standard deviation. Results were analyzed using at test and analysis of variance (ANOVA). Significance was considered at P< 0.05 using StatBox®, Version 6.0.4 (Grimmer Soft, Fr.) (2009). The tests were performed in triplicate. Results and discussion Essential oil chemical composition Seventeen compounds of the essential oil of Thymus vulgaris representing 99.24 % of the total detected constituents were identified (table 1). Among the mentioned phenolic compounds, thymol was present in the largest amounts (76.69 %) followed by ρ-cymene (9.89 %), γ-terpinene (1.92 %); caryophyllene oxide (1.69 %); β-linalool (1.51 %) and β-caryophyllen (1.32 %). A similar result was previously described by Kolsum et al. (2017) that showed 32 compounds from Thymus vulgaris with dominance of thymol (32.67 %) and p-cymene (16.68 %).The major components of the oil were thymol (36.81 %) and p -cymene (30.9 %) (Moghaddam and Mehdizadeh, 2020). Activity of essential oil on mycelial growth The results of antifungal activity assays showed that the 1 % (v/v) thyme essential oil significantly inhibited (P<0.01) the mycelium growth of Fusarium oxysporum f. sp. radicis-lycopersici after 24 h of incubation at 25 ° C compared to untreated (figure 1a). The essential oil reached an inhibition rate of 43 % for Fusarium oxysporum f. sp. radicis-lycopersici after culturing for 48 h. In the research carried out by Galovičová et al. (2021), amounts of free thyme oil between 62.5 and 500 μL.mL -1 were necessary to inhibit 46- 87 % the growth of Serratia marcescens . Thymus vulgaris essential oil completely suppressed the colony growth of Fusarium oxysporum f. sp. radicis - lycopersici (Kumar et al ., 2007; Aksit et al ., 2022). After 96 h, the mycelium growth of the strain on the essential oil-treated groups became larger, and the inhibition rate declined (10.86 %), which may be because the oil had evaporated. This scientific publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818. Rev. Fac. Agron. (LUZ). 2022, 39(3): e223941. July - September. ISSN 2477-9407. 4-6 | Table 1. Chemical constituents of Thymus vulgaris leaves essential oil using gas chromatography-mass spectrometry (GC/MS). The opposite result was observed for essential oil from Thymus. In which conidia production of Fusarium oxysporum f. sp. radicis- lycopersici was stimulated (figure 2). The average number of conidia on unamended (control) PDA media plates after 14 days of incubation at 25 ° C were 5.65x10 5 conidia.mL -1 while grown colonies amended with Thymus essential oil, production of conidia was significantly increased than the control, and up to 13.7x10 5 conidia.mL -1 . Thymol and p -cymene are the major compounds found in Thymus vulgaris and have shown potent fungicidal and/ or fungistatic activities against various phytopathogenic fungi such as Fusarium spp., Aspergillu s spp., Cladosporium spp., Mucor spp. and Rhizopus spp. (Kumar et al ., 2007). Thymol and p -cymene provide irreversible damage to the cell wall, cytoplasm membrane and nuclear membrane of filamentous fungi (Rasooliand Owlia, 2005). The effects of essential oil on the spore germination of Fusarium oxysporum f. sp. radicis-lycopersici are shown in figure 3. In the absence of NaCl, thyme oil was found to be the most effective inhibition (86 %) of spore germination. The essential oil may inhibit the cell growth and proliferation by interrupting ergosterol biosynthesis (Gao et al ., 2016). Essential oil enables us to integrate into the lipids of the cell membrane, increasing permeability occur as a due of loss of ions and reduction of membrane potential, collapse of the proton pump and depletion of the ATP pool, which eventually lead to leaking of intracellular constituents, coagulation of cell contents, lysis and cell death (Turgis et al ., 2012). Antimicrobial activity was due to disturbance of the permeability of the membrane and the release of some inhibition enzymes such as ATPase, histidine decarboxylase, and amylase from the cellular content (Scollard et al ., 2016). Carvacroland thymol disrupted the ergosterol biosynthesis and membrane integrity (Ahmad et al ., 2011). 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(c) Combined effect of salinity levels and essential oil. Figure 1. Effect of salinity levels and essential oil on mycelial growth of Fusarium oxysporum at 24, 48, 72 and 96 h. (1a) Effect of fungal load of essential oil. (1b) Effect of salinity on mycelial growth. (1c) Effect of salinity and essential oil on mycelial growth. ns, *, ** and *** non- significant, significant at 0.05, 0.01 and 0.001 probability level, respectively. Compounds RT.min- 1 RI Content (%) Thujene α-Pinene cis-Sabinene hydrate Camphene β-Myrcene α-Terpinene β- Phelladrene γ-Terpinene ρ-Cymene Carvacryl Acetate β-Linalool 4-Terpineol β-Caryophyllen Borneol Caryophyllene oxide Thymol Eugenol 10.24 10.32 10.66 10.76 11.74 11.88 12.18 12.55 12.80 13.72 15.36 16.07 16.12 17.19 20.18 21.99 21.61 1140 1165 777 846 889 916 1012 965 1024 1303 998 1107 1319 1012 1479 1196 1300 0.71 0.49 0.46 0.14 0.71 0.58 0.37 1.92 9.89 1.19 1.51 0.65 1.32 0.49 1.69 76.96 0.16 Total identified (%) 99.24 (a) Effect of fungal load of essential oil iVBORw0KGgoAAAANSUhEUgAAAoAAAAFqCAYAAACK+utaAAA8oUlEQVR42u2dW3BV1fnADXfk kiAJN2lE5BrLVENHual50KcaJ6gFBTtlOqOvjlQQmIrUdsp0+tCpF0QgeOlk0oe+oIBQ0anN eKGdVqu2o4BBtCi3gNwSIJH1n2/Nfx13dvb1nH3O2Zffb2aPkrOTc/Z39lnrd7611reuUAAA AACQKa4gBAAAAAAIIAAAAAAggAAAAACAAAIAAAAAAggAAAAACCAAAAAAIIAAAAAAgAACAAAA AAIIAAAAAAggAAAAACCAAAAAAIAAAgAAAAACCNmgu7tbNTY2qiuuuKLX8cQTT5Tt9TzzzDNF 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Effect of salinity and essential oil on conidia production by Fusarium oxysporum. Values with different letters show significant difference (P≤0.05). This scientific publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818. Boumaaza et al. Rev. Fac. Agron. (LUZ). 2022, 39(3): e223941 5-6 | Effects of NaCl on mycelial growth, germination and conidia production After 24 h of incubation, the mycelial growth eached a diameter of 0.82 mm in the control. Whereas under gradual concentrations of NaCl, colony diameters were 1.2 mm at 0.3 % NaCl, 1.27 mm at 0.6 %, 1.3 mm at 1 %, 1.35 mm at 1.5 % and 1.2 mm at 2 % respectively (figure 1b). The average mycelial growth was significantly in the saline medium over the five levels of NaCl compared to the control. Figure 1b shows that the saline medium has a significant influence on the stimulation of mycelial growth compared to the control. Similar supporting results have been reported (Boumaaza et al. , 2015). Figure 2 shows that the application of sodium salt caused a significant increase in the conidia production at various concentrations tested compared with control (P< 0.001). By adding 2 % of NaCl to the culture medium, an increase in spore production by 13.6x10 5 conidia.mL -1 can be obtained compared to the control 5.65x10 5 conidia.mL -1 . Ahigh concentration of sodium chloride showed to be the most favorable to the sporulation. The germination of conidia that were affected by higher levels of NaCl is shown in figure 3. An inhibition of 2.66 % in germination was observed at 2 % salinity. However, the lower levels of NaCl (0.3, 0.6 and 1 %) were significantly not active against Fusarium oxysporum f. sp. radicis-lycopersici . Figure 3. Effect of salinity and essential oil on conidia germination by Fusarium oxysporum. Values with different letters show significant differences (P≤0.05). Combined effects of salinity and essential oil on Fusarium oxysporum The combined effects of salinity levels and essential oil on Fusarium oxysporum are shown in figure 1c. After 48h of incubation, the growth of the mycelium treated with essential oil decreased to higher salinity levels. Compared to untreated salinity (3.4 mm), the mycelium growth achieved 3.63, 3.68, 3.26, 3.41 and 2.48 mm was obtained by the concentrations of 0.3, 0.6, 1, 1.5 and 2 % NaCl, respectively. The results shown in figure 2 revealed that the combined effects of salinity levels and essential oil caused a significant increase in spore production at various levels tested compared with control. However, this combination increase the antifungal activity of essential oil compared to oil alone (13.7x10 5 conidia.mL -1 ), the conidia production achieved 2.66, 3.5, 3.6, 5.2 and 9.76x10 5 conidia.mL -1 were obtained by with concentrations of NaCl 0.3, 0.6, 1, 1.5 and 2 %, respectively. 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The spore germination achieved of 20.66, 57, 58.66 and 58 % was obtained by the concentrations of 0.3, 0.6, 1 and 1.5 % respectively. The results of this study indicated that salinity decreases the efficacy of essential oils against the pathogen. According to Perumal et al . (2016), the efficacy of essential oils against the microorganisms mainly depend on the characteristics of their components and influenced by many factors such as high temperature, low water activity, low nutrient conditions and UV light for effective establishment and disease control. Several studies have investigated the control efficacy of essential oils against Fusarium sp. (Xing et al., 2014). To date, there have been limited studies on the use essential oils in mycology under salinity conditions. In accordance with Biswas et al . (2011), essential oil contents are also affected by environmental factors such as salinity, water stress and soil pollution (Stancheva et al ., 2014). 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