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Marziyeh Farsia, Farzin Abdollahib, Amin Salehic, Shiva Ghasemid 1-Former MSc Student of Horticultural Science Department, Faculty of Agriculture and Natural Resources, University of Hormozgan. HYPERLINK [email protected] [email protected] 2-Assistant Professor of Horticultural Science Department, Faculty of Agriculture and Natural Resources, University of Hormozgan (Correspondence Author). HYPERLINK [email protected] [email protected] 3- Associate Professor of Department of Agronomy and Plant Breeding, Faculty of Agriculture, Yasuj University. HYPERLINK [email protected] [email protected] 4- Instructor of Horticulture Crops Research Department, Qazvin Agricultural and Natural Resources Research and Education Center, Agricultural Research, Education and Extension Organization (AREEO), Qazvin, Iran. HYPERLINK [email protected] [email protected] Declarations of interest none. Abstract Key words Essential oil yield and percent, monoterpene hydrocarbons, oxygenated monoterpenes, water stress 1. Introduction Medicinal and aromatic plants are important in terms of their flavors and medicinal and pharmaceutical properties, along with human consumption, feeding animals and ornamental uses (De Feo et al., 2005). A large number of these species belong to the family Lamiaceae, and within this family, marjoram (Origanum majorana L.) is economically important (Omidbaigi and Bastan, 2005). Marjoram grows wildly in the region of the Mediterranean Sea, India and Middle East. It is also cultivated in gardens and fields in many European, Asian and North American countries for consumption and pharmaceutical purposes (Nowak and Ogonowski, 2010). The oil is used in perfumery for its spicy herbaceous notes (Vera and Chang-Ming, 1999). The buds and flowers are oil-richest, leaves contain less, and the stems carry only trace amounts (Nowak and Ogonowski, 2010). The composition of oils from various marjoram species has previously been investigated (Novak and Ogonowski, 2010 Omidbaigi and Bastan, 2005 Vera and Chang-Ming, 1999 Hornok, 1993). The species is frequently used as a carminative, expectorant, stimulant, tonic, antiviral, and antifungal agent among other medical applications (Charles 2012 Mozafarian, 2012). Although different chemical compounds have been isolated from wild marjoram (Origanum vulgare L.), only the essential oil is considered the most commercially important natural product, displaying the main biological activities (Morshedloo et al., 2017). Drought is the main abiotic stressor around the world and drastically reduces crop yields by affecting plant morphology, physiology and biochemistry. It destructively influences the capability to meet food demands of an ever-increasing global population (Tester and Langridge, 2010). Water deficit in plants may cause physiological disorders, such as a reduction in photosynthesis and transpiration (Sarker et al., 2005), and there are many studies in the case of medicinal plants that show water stress may cause significant changes in the yield and composition of essential oils. For example, water deficit decreases the oil yield and changes the oil composition of wild marjoram, thymus (Thymus carmanicus), Parsley (Petroselinum crispum Mill. Nym. ex A.W. Hill), rosemary (Rosmarinus officinalis L) and anise (Pimpinella anisum L.) (Morshedloo et al., 2017 Bahreininejad et al., 2014 Petropoulos et al., 2008 Singh and Ramesh, 2000 Zehtab-Salmasi et al., 2001). In recent years, scientists have been looking for new alternatives to conventional methods for plant protection, and simultaneously for improvements in health properties and bioactive compound content. One of these methods is elicitation, which is induction of natural plant resistance mechanisms using biotic or abiotic factors. Elicitation can be an important strategy towards obtaining improved production of plant secondary metabolites in vivo (Hussain et al., 2012). It has been reported that plant hormones such as jasmonic acid and methyl jasmonic acid (MeJA), are potent elicitors and/or signaling agents and play a key role in plant growth and development they are also involved in stress responses in plants (Zhao et al., 2005). Some studies have suggested that exogenous application of these compounds under normal and stressful conditions improved the levels of phytochemicals and secondary metabolites in some crops, including medicinal crops (Andrys et al., 2017 Ghasemi Pirbalouti et al., 2014 Hamedi et al., 2014 Kim et al., 2006 Malekpoor et al., 2015 Miranshahi and Sayyari, 2016 Zhao et al., 2005 Zlotek et al., 2014). Since the effects of MeJA on essential oil content and composition of marjoram (Origanum majorana L.) have not yet been reported under water stress conditions, the objective of this study was to evaluate the effects of two different MeJA concentrations on essential oil content and responses of oil composition changes under three water deficit conditions. 2. Materials and Methods Study site and experimental design This study was conducted at the research greenhouse of Faculty of Agriculture and Natural Resources of University of Hormozgan, Bandar Abbas, Iran, during the period between 2014 and 2015. The marjoram seeds were provided from seed gene bank at the Forest and Rangeland Research Institute in Tehran, Iran. Twenty-five uniform marjoram seed sets were sown on 12 November 2014 in 6 L plastic pots filled with a thoroughly mixed 211 ratio of sieved field soil, sand, and leaf mold. The pots were kept in a greenhouse at a temperature varying between 25 and 27C during the day and between 18 and 22C during the night and photoperiod varying between 12 and 14 h in days, 10 and 12 h at night. The relative humidity of the greenhouse was kept constant by about 40. Gravimetric soil water content at field capacity (FC) and permanent wilting point (PWP) that estimated with pressure plate apparatus were 19.1 and 4.4 respectively. Fourteen days after seedling emergence, plants were thinned and 10 vigorous plants remained in each pot. 2.2 Plant weight, essential oil yield and percentage measurement At 25 February 2015, in the flowering stage (10 days after forth spray of MeJA), total plants of each pot were removed. After determination of fresh weight, plant samples were air-dried at room temperature and then total shoot (above ground) dry weight and essential oil percentage (EO) were measured. Fresh and dry weight were measured as weight per plant. EO was determined on a dry weight basis from 50 g of air-dried herb harvested from each pot. Air-dried shoot of the marjoram were subjected to hydrodistillation for 3 h with a Clevenger-type apparatus according to the procedures of Kulisic et al. (2004). To calculate the EO yield (mg plant-1), multiplication of EO percent in plant dry weight was used. In order to perform essential oil composition analysis, the oil was dried over anhydrous sodium sulfate and stored in sealed vial at -18 C before analysis. 2.3 Determination of essential oil composition 2.4 Statistical analysis A one-way analysis of variance (ANOVA) was performed between treatment samples in a completely randomized design in three replications. The data were subjected to analysis of variance at 5 level of significance using SAS software (SAS Institute, 2003). Significant levels of differences for all measured traits were calculated and means were compared by the LSD test at P 0.05. 3. Results and Discussions 3.1 Effect of water deficit on plant weight, essential oil percentage and essential oil yield Severe water deficit significantly decreased fresh and dry weight of marjoram 14.4 and 15.1, respectively, when compared with the control treatment. However, moderate water deficit had no significant effect on these traits (Table 1). Similar to these results, some studies indicated that water deficit reduced plant growth and biomass in oregano (Oreganum vulgare) (Morshedloo et al., 2017 Said-Al Ahl and Hussein, 2010). It seems that biomass reduction under water deficit conditions is as adaptive strategy to limit the effects of water stress on plant cells and survival under stress conditions (Yadav et al., 2014 Farooq et al., 2009). Furthermore, under water deficit conditions the biomass reduction may be due to a decrease in photosynthesis (Shao et al., 2008), lower availability of sufficient moisture around the root and, therefore, lower water and nutrient absorption, leading to decreased plant biomass (Singh et al., 1997) The EO of marjoram increased significantly about 9.18 and 10.20 under moderate and severe water deficit, respectively, compared to control However, there was no significant difference between the water conditions (Table 1). In confirming these results, Morshedlo et al. (2017), Miranshahi and Sayyari (2016), and Bahreininejad et al. (2014) reported that water stress caused significant increase in EO in oregano, summer savory and thyme, respectively. Stressful environments can stimulate essential oil production (Sangwan et al., 2001). This stimulation of essential oil production might be due to increase in density of the glandular trichomes producing essential oil on the leaf tissue, resulting in the increase in amount of essential oil accumulation per unit of leaf tissue (Delfine et al., 2005). The highest (20.3 mg plant-1) and lowest (17.1 mg plant-1) EO yield were obtained in moderate and severe water stress condition, respectively (Table 1). Moderate water deficit level has significantly higher EO yield than severe water deficit (18.3). This may be because of higher dry weight and EO in moderate water deficit in comparison with severe water deficit (Table 1). In agreement with our results, Shokrani et al. (2012) and Baher et al. (2002) reported an increase in essential oil content and yield in calendula (Calendula officinalis) and savory (Satureja hortensis) with increasing water stress severity and higher yield of essential oil obtained from mild water stress, respectively. In contrast with these results, Bahreininejad et al. (2014) showed that water deficit significantly decreased oil yield about 43.0 in thyme. Furthermore, some studies indicate that irrigation regime had no significant effect on EO yield in aromatic plants (Jordan et al., 2003 Khazaie et al., 2008). In general, moisture deficit increases the essential oil content of more medicinal and aromatic plants, because in case of stress, more metabolites are produced in the plants and substances prevent oxidization in the cells. Essential oil helps plant to adopt easier to the environmental stress conditions (Turtola et al., 2003). Table 1. Effects of water deficit on fresh and dry weight, essential oil percentage and yield of marjoram plants a. Water deficit Fresh weight (g plant-1)Dry weight (g plant-1) EO EO yield (mg plant-1) Control b7.87 a1.86 a0.98 b18.2 bModerate7.75 a1.90 a1.07 a20.3 aSevere6.74 b1.58 b1.08 a17.1 bLSD1.000.240.081.5a In each column, the means with at least one similar letter do not differ significantly (P 0.05). b Control, moderate and severe, represent pots irrigated when gravimetric soil water contents were 100, 75 and 50 FC, respectively. 3.2 Effect of MeJA on plant weight, essential oil percentage and essential oil yield This study indicated that MeJA did not have a significant effect on fresh and dry weight of marjoram. However, it should be noted that the use of MeJA caused a negative effect and decrease in these traits slightly (Table 2). Contrary to these results, Zlotek et al., 2016 and Miranshahi and Sayyari (2016) reported that there was positive effect of the jasmonic acid (JA) and MeJA elicitation on the growth and yield parameters of the thyme (Ocimum basilicum L.) and summer savory (Satureja hortensis), respectively. Similar to our results, some researchers reported that foliar application of jasmonic acid (JA) had no significant effects on thyme (Zlotek et al., 2016 Alavi-Samani et al., 2015) and st. Johns Wort (Hypericum perforatum L.) growth parameters and plant growth indices (Hamedi et al., 2014). It is concluded that the influence of JA elicitation on plant growth may depend on plant condition, the elicitor type and dose (Zlotek et al., 2016). Table 2. Effects of MeJA on fresh and dry weight, essential oil percentage and yield of marjoram plants a. MeJA 3.3 Effect of water deficit and MeJA interaction on plant weight, essential oil percentage and essential oil yield In most cases, MeJA decreased EO and EO yield (Table 3). This negative effect was more obvious under moderate water deficit, so that under this condition, application of MeJA causes significant reductions in EO and EO yield of about 16.9 and 12.0, respectively (Table 3). These results indicate that under moderate stress condition, although there was elicitation effect of MeJA on the fresh and dry weight, because EO decreased significantly EO yield also decreased significantly. Contrary to our results, some studies have reported that with application of MeJA or JA under non-stress and stress conditions, EO and EO yield of some medicinal and aromatic plants increased (Andrys et al., 2017 Miranshahi and sayyari, 2016 Zlotek et al., 2016 Malekpoor et al., 2015). The essential oil production depends not only on genetic factors, but also on environmental factors. Plant growth regulators can change essential oil content through affecting plant growth, essential oil biosynthesis, and the number of oil storage structures. On the other hand, the time of exogenous application and kind of growth regulators can affect essential oil content (Sharafzadeh, 2011). Table 3. Effects of water deficit and MeJA interaction on fresh and dry weight, essential oil percentage and yield of marjoram plants a. Water deficit MeJA b Control, moderate and severe, represent pots irrigated when gravimetric soil water contents were 100, 75 and 50 FC, respectively. 3.4 Marjoram essential oil composition 3.5 Effect of water deficit on chemical compositions of marjoram essential oil 3.6 Effect of water deficit and MeJA interaction on chemical compositions of marjoram essential oil Table 5. Effects of water deficit on essential oil composition of marjorama. b Control, moderate and severe, represent pots irrigated when gravimetric soil water contents were 100, 75 and 50 FC, respectively. Table 6. Effects of MeJA on essential oil composition of marjorama. b Control, moderate and severe, represent pots irrigated when gravimetric soil water contents were 100, 75 and 50 FC, respectively. 4. Conclusion Nowadays, drought stress significantly decreased crops yield in Iran. Therefore, to decrease hazardous effects of drought stress recently new approaches such as cultivation of crop species, including medicinal plants, with low water requirements and application of strategies (including plant hormones like MeJA application as an elicitor) that improve their physiological traits and change EO composition in water deficit conditions are mentioned. Results of our study indicated that severe water deficit decreased plant fresh and dry weight and essential oil (EO) yield of marjoram. When water deficit level increased moderately, EO production increased so that maximum EO and yield were observed in moderate water deficit and under severe water deficit conditions these traits decreased significantly. Although water deficit could not change marjoram essential oil components, but when MeJA was applied marjoram essential oil composition changed. So that with increasing in water deficit level, MeJA could decrease monoterpene hydrocarbons components content, however, under this condition oxygenated monoterpenes increased. As a result, when MeJA was applied under severe water deficit, signs of drought stress were limited. This may be due to elicitor role of MeJA in stimulating oxygenated monoterpenes production and, therefore, improved essential oil production under water stress conditions. 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