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Profiling of Essential Oils Components and Polyphenols for Their Antioxidant Activity of Medicinal and Aromatic Plants Grown in Different Environmental Conditions

Antonios Chrysargyris
Maria Mikallou
Spyridon Petropoulos
2 and
Nikolaos Tzortzakis
Department of Agricultural Sciences, Biotechnology and Food Science, Cyprus University of Technology, 3603 Limassol, Cyprus
Department of Agriculture, Crop Production and Rural Environment, University of Thessaly, N. Ionia, 38446 Magnissia, Greece
Authors to whom correspondence should be addressed.
Agronomy 2020, 10(5), 727;
Submission received: 27 April 2020 / Revised: 13 May 2020 / Accepted: 15 May 2020 / Published: 19 May 2020


In the present study, the yield, the chemical composition, and the antioxidant activities of the essential oils (EOs) of eight medicinal and aromatic plants (MAPs) cultivated under two environmental conditions characterized by a different altitude (namely mountainous and plain) were evaluated. Cultivation at different environmental conditions resulted in significant differences in the chemical composition and antioxidant activity for most of the studied species. In particular, high altitudes resulted in increased phenolic compounds’ content and antioxidant activity for artemisia plants, while specific parameters increased in the case of spearmint (total phenols) and rosemary (flavonoids). In contrast, in pelargonium, all the tested parameters were positively affected in the plain area, whereas, for laurel and sage, only flavanols remained unaffected. EO yield in mountainous pelargonium and spearmint decreased while, in mountainous laurel, pelargonium and spearmint increased when compared to plain areas. In addition, the major EO constituents’ content for most of the species were affected by environmental conditions. The 2,2-diphenyl-1-picryl-hydrazyl-hydrate (DPPH) and ferric reducing antioxidant power (FRAP) were variably correlated with total phenols, flavonoids, and flavanols, depending on the species and the altitude. Lastly, in limited cases, antioxidant activity (DPPH or FRAP values) was positively correlated with some EO components (e.g., borneol and β-pinene in artemisia and laurel plants grown in the plain, respectively, or 1,8-cineole in mountainous grown verbena plants). In conclusion, environmental conditions (altitude) affected antioxidants’ content and EO yield and composition of the studied MAPs. These findings can be used to introduce cultivation of MAPs in specific ecosystems for the production of high added value products.

1. Introduction

Medicinal and aromatic plants (MAPs), also known as herbs or spices, and their relevant plant extracts and essential oils (EOs) have been highly appreciated and widely used for centuries despite the lack of scientific evidence for their actual bioactive mechanisms and functions, which are still under investigation [1,2,3]. Contemporary dietary patterns also prescribe such MAPs as functional foods, i.e., foods that offer additional physiological benefits beyond the usual nutritional requirements such as preventing or delaying the onset of chronic diseases [4]. The global interest in MAPS is reflected in the trade of MAPs as raw material, which is approximately 440,000 tons per year at a total value of $1.3 billion USD. A total of 25% of this monetary value is marketed in Europe [5].
Food products rich in antioxidants are well appreciated since they can act as scavengers of reactive oxygen species (ROS) and also help decrease the impact of age-related chronic diseases [6]. Therefore, MAPs have been the focus of scientific research and the food and pharmaceutical industry due to their well acknowledged antioxidant capacity [3,7]. A diverse range of secondary metabolites such as phenolic compounds are biosynthesized by plants as part of their protection mechanism toward oxidative damage by ROS and abiotic and biotic stressors, while these compounds may also have protective effects on humans when MAP and/or their components are ingested through diet [8]. The antioxidant capacity of phenolic compounds involves a combination of different mechanisms including free radical scavenging, donation of hydrogen atoms, single oxygen quenching, metal ion chelation, and activities as an oxidation substrate [8].
The Mediterranean basin is abundant in MAPs with more than 10,000 species being identified so far, which have been widely used in the Mediterranean diet [9,10,11]. The important bioactive properties of these species have been systematically reported in ethnobotanical and ethnopharmacological studies [12,13,14,15,16,17]. However, further investigation is needed to reveal and define their precise pharmaceutical and functional properties as food additives and novel antioxidants [18]. Biological activity and phytochemicals of MAPs show a great variability depending on the cultivation area, the climatic conditions, and the genetic material [19,20,21]. According to Kofidis and Bosabalidis [22], altitude was suggested to be one of the most important ecological factors affecting bioactivities of MAPs as specific environmental factors such as light and wind intensity, mean temperature, ozone levels, and partial CO2 pressure, which may vary between different altitudes. Moreover, MAPs bioactive properties are often associated with the presence of secondary metabolites with antioxidant potential such as phenolic compounds [23,24]. However, special attention should be given prior to recommending the use of MAPs in human diet, since, in several occasions, the intake of high doses of secondary metabolites and potentially harmful substances (e.g., heavy metals and anti-nutritional factors) may cause severe toxicity and adverse health effects [25,26,27,28]. Therefore, further research is needed to evaluate possible toxic effects and establish recommended daily allowance (RDA) levels, especially for people with medical conditions [29,30,31].
Apart from the use of MAPs as herbs and decoctions, their EOs have also found very important uses in the food and pharmaceutical industry [32]. Several studies with EOs revealed significant antioxidant [33,34] and antimicrobial properties [35,36,37,38] and further increased the interest to use EOs as natural antioxidants and antimicrobial agents instead of synthetic compounds, which are currently receiving criticism due to harmful effects on human health [36,38,39]. However, despite the increased interest and the great number of MAPs throughout the world, only approximately 10% of the already known EOs have received attention due to their varied biological activities [2,3]. Currently, they are widely used in the food, cosmetics, and pharmaceutical industry [40,41].
MAPs cultivation in Cyprus shows promising prospects as crops have low requirements in agrochemicals, irrigation water, man power, and energy [42]. They also exhibit tolerance to arduous climatic conditions such as high temperatures, winds, and drought [43,44]. All these important features could help the sustainable development of rural areas and also reduce the threats arising from wild harvesting of MAPs [45]. Although the island’s soil and climatic conditions are ideal for MAPs growth, their cultivation is not yet widespread because of limited availability of agricultural land due to other uses, i.e., tourism and constructions. Based on the above, it is recommended to evaluate potential areas and/or cultivation practices that may provide high quality and added value products of MAPs [46], so that farmers could shift to these crops and establish economically viable farms. Due to the increased global demands of high value MAPs, Cyprus with a long history on MAPs’ cultivation and uses could become a significant spot for producing and exporting high quality raw materials of MAPs to other countries that are more industrially developed for further processing.
So far, several studies examined the correlation of total phenolics and/or phenolic compounds’ content with the antioxidant activity of various MAPs products such as infusions, decoctions, and EOs [24,47,48,49,50]. For example, the antioxidant activity and phenolics compounds in 10 selected MAPs from Serbia, revealed a positive correlation of phenolics and tannins, but also a proportional increase of antioxidants with total phenolics increase [50]. However, the correlations of the main compounds of EOs with the phenolic compounds content and the antioxidant activity of leaves are scarcely explored. Therefore, in an attempt to contribute to the existing knowledge, the aim of this work was to compare medicinal and aromatic plants grown in Cyprus at different environmental conditions (altitudes: mountainous and plain areas) in view of revealing possible correlations between their leaf antioxidant activity and their essential oil yield and composition. The selection of the studied plant species was based on their popularity and their recommendation for use.

2. Materials and Methods

2.1. Plant Material and Growing Conditions

The studied medicinal and aromatic plants were as follows: artemisia (Artemisia abrotanum L.), pelargonium (Pelargonium roseum L.), laurel (Laurus nobilis L.), rosemary (Rosmarinus officinalis L.), spearmint (Mentha spicata L.), lavender (Lavandula angustifolia L.), lemon verbena (Aloysia triphylla L.), sage (Salvia officinalis L.), and their parts are presented in Table 1. The plants were identified by staff members of the Cypriot National Agricultural Department.
In the current study, two areas with different environmental conditions (for simplicity, the term altitude will be used, considering the differences in the microclimates as described by Kofidis and Bosabalidis [22]) were selected, the mountainous area of ″Agros″ village (34°53’44.10″ N; 33°01’13.86″ E) and the plain area of Nicosia (“Athalassa”; 35°08’07.65″ N; 33°24’07.22″ E). The village area of ″Agros″ is located at 880 m above sea level. The climate is dry with low temperatures and snow precipitation during winter. The soil has sand-silk texture. On the other hand, the plain area of ″Athalassa″ is located 141 m above sea level with mild winter and a dry-hot summer. Detailed climatic conditions of the selected areas are described in supplementary material (Table S1).
Plant species from the plain area harvested from the farm of the Cypriot National Agricultural Department were 2–5 years old (except for laurel where plants were approximately 15 years old). Common cultivation practices were applied and plants were frequently irrigated (~weekly/biweekly during the growing period) and common fertilizers were applied (20-10-10 (N-P-K) once a year in base dressing and 19-19-19 (N-P-K) every second month in side dressing. Mountain species harvested from public green areas/parks of ″Agros″ village were of a different age and were grown under non-commercial cultivation practices, which means they received conservation practices (periodical irrigation and fertilizer application). Plants for public green area use and landscaping normally originated from the Cypriot National Agricultural Department or from nurseries that collaborate with this department. Plant tissues (six samples/area/species) of the above ground parts (leaves or leaves and stems) (see Table 1) were collected early to mid-October and transferred within an hour to the laboratory. Each sample was divided into two batch samples. One batch was air-dried at room temperature for approximately 7 days and used for the essential oil extraction (see Section 2.4) while the other batch was stored at −20 °C for the chemical analyses described in Section 2.2 and Section 2.3.

2.2. Polyphenol Extraction and Analyses

2.2.1. Extract Preparation

Six samples (0.5 g) of freshly cut plants (pooled by two individual plants/sample) from each treatment were milled with 10 mL methanol (80%) [59]. The extracts were centrifuged for 30 min at 4000× g at 4 °C (Sigma 3-18K, Sigma Laboratory Centrifuge, Germany). After centrifugation, the supernatant was transferred to a 15-mL falcon tube, and stored at 4 °C until further analyses (within 24 h) for evaluating total phenolics, flavonoids, and flavanols content and total antioxidant activity.

2.2.2. Total Phenolics

The total phenolic compounds content of the methanolic extracts was determined by using the Folin–Ciocalteu reagent (Merck), according to the procedure described by Tzortzakis et al. [59]. A total of 125 μL of plant extracts were mixed with 125 μL of the Folin–Ciocalteu reagent. The mixture was shaken before the addition of 1.25 mL of 7% Na2CO3, adjusted with distilled water to a final volume of 3 mL, and mixed thoroughly. After incubation in the dark for 90 min, the absorbance of extracts at 755 nm was measured in comparison to the prepared blank. Total phenolic compounds content was expressed as μmol of gallic acid equivalents per gram of fresh weight (μmol GAE g−1 fw) through a calibration curve prepared with gallic acid. All samples were analysed in triplicate.

2.2.3. Total Flavonoids and Flavanols

The total flavonoid content was determined according to the aluminium chloride colorimetric method [60]. Plant extracts and 0.75 mL of 5% sodium nitrite (NaNO2) were incubated for 6 min. After the incubation, 0.15 mL of AlCl3 solution (10%) was added. After an additional time of 5 min, 0.5 mL of NaOH (1 M) solution was added and the final volume was adjusted to 2.5 mL with the addition of distilled water. The solution was mixed thoroughly, and the absorbance was measured at 510 nm. The total flavonoids content was expressed as rutin equivalents (mg rutin g−1 fw).
Total flavanols content was determined according to Tabart et al. [61]. In more details, 1 mL of catechin solution (0–300 μg mL−1 in methanol) or test solution (150–250 μg mL−1 polyphenols in methanol) were added in test tubes. Then, 2.5 mL of methanol (control) or 1% vanillin solution in methanol and 2.5 mL of 9 M HCl in methanol (test samples) were added. The reaction mixture was incubated for 20 min at 30 °C and the absorbance at 500 nm was measured. The catechin solution was used for preparing the calibration curve. Results were expressed as catechin equivalents (mg catechin g−1 fw).

2.3. Antioxidant and Reducing Activity

The antioxidant and reducing activity of plant extracts was evaluated using the 2,2-diphenyl-1-picryl-hydrazyl-hydrate (DPPH) and ferric reducing antioxidant power (FRAP) assays according to Wojdyło et al. [62] with some modifications [63]. The DPPH radical scavenging activity of plant extracts was measured from the bleaching of the purple-colored 0.3 mM solution of DPPH, which consisted of 1 mL of the DPPH solution, 1.98 mL of 50% methanol, and 0.02 mL of plant extract. After shaking, the mixture was incubated at room temperature in the dark for 30 min, and then the absorbance was measured at 517 nm. DPPH radical-scavenging activity was expressed as the inhibition percentage (I %) and was calculated using the following formula.
DPPH radical scavenging activity I (%) = [100 − 100 × (Abs − Abb)/Abc]
where Abb is the absorbance of the blank sample, Abs is the absorbance of the test sample, and Abc is the absorbance of the control with DPPH and 50% methanol.
For the ferric reducing/antioxidant power (FRAP) assay, a sample of 3 mL of freshly prepared FRAP solution (0.3 mol L−1 acetate buffer, pH 3.6), containing 10 mmol L−1 TPTZ (Tripyridil-s-triazine) and 40 mmol L−1 FeCl3·10H2O and 20 μL of extract (50 mg mL−1) were incubated at 37 °C for 4 min and the absorbance was measured at 593 nm. The changes in absorbance were then converted into a FRAP value by relating the change of absorbance at 593 nm of the test sample to that of the standard solution of trolox ((±)-6-Hydroxy-2,5,7,8-tetramethylchromane-2-carboxylic acid). The results were expressed as mg trolox g−1 fw.

2.4. Essential Oil Extraction and Gas Chromatography/Mass Spectrometry Analysis

Extracting essential oils was carried out according to the protocol previously described by the authors [64]. Aerial parts were collected and air-dried at room temperature. The dried tissues (40–50 g for each treatment) were used for the essential oil extraction using a Clevenger apparatus. Each extraction lasted for 3 h, while each treatment was replicated three times. The essential oil (dried over anhydrous sodium sulphate) yield was measured and calculated as percentage of oil per dry weight (dw) [63]. The obtained essential oils were kept in amber glass bottles at −20 °C until GC/MS analysis was performed.
Analytical gas chromatography was carried out with a Shimadzu GC2010 gas chromatograph interfaced Shimadzu GC/MS QP2010 plus mass spectrometer based on the protocol previously described by the authors [63]. An aliquot of 2 μL of each sample was injected in a split mode (split ratio 20:1) into the gas chromatograph fitted with a ZB-5 column (Zebron, Phenomenex, Torrance, CA, USA) coated with 5% pheny to 95% dimethylpolysiloxane with film thickness of 0.25 μm, length of 30.0 m, and a diameter of 0.25 mm. The flow of the carrier gas (helium) was 1.03 mL min−1. The injector temperature was set to 230 °C. Electron impact mass spectra with ionization energy of 70 eV was recorded at the 35–400 m z−1. The column temperature was programmed to rise from 60 °C to 240 °C at a rate of 5 °C min−1 with a 5 min hold at 240 °C. The solution of standard alkanes mixture (C8–C20) was also analyzed using the above conditions.
Components were identified by comparing their retention indices (RI) relative to n-alkanes (C8–C20) with those of the literature or with those of authentic compounds when available. Further identification of compounds was carried out by matching the recorded mass spectra with those stored in the NIST08 mass spectral library of the GC–MS data system and published mass spectra in the literature [63]. The percentage of individual compounds was based on peak area normalization without using correction factors.

2.5. Statistical Methods

A two factor (Species and Environmental Conditions, namely Altitude) factorial experiment was carried out. The statistical treatment of the results was carried out using a two-way analysis of variance (ANOVA) by using the IBM SPSS v.22 software for Windows. The Student’s t-test (p < 0.05) was used for the comparison of means when the effect of altitude was significant, while the Duncan Multiple Range Test was used for comparing means in the cases where the effect of species and the interaction of species × altitude were significant. Mean values are presented as treatment mean ± SE of six biological measurements (n = 6) for antioxidants and for three biological measurements (n = 3) for essential oils analysis. The correlation coefficients between mountainous and plain species and their antioxidant capacity and essential oil components were also determined.

3. Results and Discussion

3.1. Total Phenols, Flavonoids, Flavanols, and Antioxidant and Reducing Activity

Phenolic compounds are one of the most important classes of natural antioxidants and are closely related with the antioxidant activity of plant tissues [49,65]. In this study, we tried to determine whether the content of phenolic compounds (total phenolics, flavonoids, and flavanols) and the antioxidant activity of eight MAP species were affected by the altitude as previously reported [48,66]. Table 2 presents the effects of environmental conditions-altitude (mountain versus plain) and species on the phenolic compounds content as well as on the antioxidant activity of the examined MAP species. The two-way ANOVA revealed a significant (p < 0.001) interaction between the tested factors (species and altitude) for all the tested parameters. Moreover, the species factor significantly (p < 0.001) affected all the tested parameters (p < 0.01), whereas altitude only affected phenolics (p < 0.01), DPPH, flavanols (p < 0.001), and EO yield (p < 0.01).
In general, altitude affected antioxidant capacity of the examined species, as plain plants presented higher flavanols and DPPH (7.16 ± 2.29 mg catechin g−1 fw and 25.02 ± 3.72 mg trolox g−1 fw, respectively) than mountainous grown plants (Table 2). When comparing all the species regardless of altitude, total phenol levels were higher in laurel and pelargonium (113.02 ± 5.31 and 112.03 ± 17.69 μmol GAE g−1 fw, respectively), whereas lemon verbena exhibited the lowest content of total phenols. The highest levels of flavonoids were found in lavender (17.31 ± 1.03 mg rutin g−1 fw) while pelargonium, rosemary, lemon verbena, and sage revealed the lowest levels of flavonoids. Moreover, pelargonium revealed the highest levels of flavanols (29.14 ± 5.78 mg catechin g-1 fw), which was followed by rosemary (8.48 ± 0.74 mg catechin g−1 fw), whereas significantly lower values were found in most of the examined species. Similarly, pelargonium was the species that revealed the higher antioxidant and reducing activity for both DPPH and FRAP assays (47.70 ± 10.41 and 25.28 ± 1.91 mg trolox g−1 fw, respectively).
On the other hand, when considering the combined effect of the tested factors, pelargonium plants grown in plain areas presented the highest content of total phenols and total flavanols, and the highest antioxidant activity for DPPH and FRAP (166.9 ± 12.0 μmol GAE g−1 fw, 47.1 ± 3.8 mg catechin g−1 fw, 80.9 ± 6.0 mg trolox g−1 fw, and 29.9 ± 2.1 mg trolox g−1 fw, respectively). Moreover, the highest levels of flavonoids (22.3 ± 1.0 mg rutin g−1 fw) were found in mountainous artemisia plants.
In previous studies, the effects of collection site on total phenolics and antioxidants has also been reported in Salvia argentea, S. officinalis, and S. verbenaca since eco-geographical characteristics may alter the biosynthetic pathways of secondary metabolites [47,67,68,69]. The effect of altitude on phenolic compounds content and the antioxidant activity has been reported in various plants species, including Thalictrum foliolosum DC. (TF) [67], Potentilla fruticosa L. [68], and Sphagnum junghuhnianum [69] among others, since several environmental factors such as elevated CO2, water availability, and differences in temperatures’ solar radiation may affect secondary metabolism and trigger the biosynthesis of bioactive compounds [70,71,72]. Antioxidant activity of plants is positively correlated with the levels of total phenolics content and, according to Žugić et al. [50], MAP species with less than 10 mg GAE g−1 of the extract (or <58.78 μmol GAE g−1) exhibited the lowest antioxidant activity. This is a finding that was also observed in our study in the case of artemisia (plain), pelargonium, sage (mountainous), and lemon verbena (both sites) (Table 2). Moreover, Pirbalouti et al. [73] reported flavonoids extracts to vary from 7.63 to 14.52 mg of rutin g−1 of tissue in Echinophora platyloba, Heracleum lasiopetalum, and Kelussia odoratissima and suggested the use of MAPs as an alternative preservative and dietary source of antioxidants in the food industry (i.e., pickles). Similarly, a relatively high total flavonoids content was observed in lavender, laurel, and artemisia in the present work under different altitudes.

3.2. Essential Oil Yield and Composition

The two-way ANOVA reveled that EOs yield was affected by altitude (p < 0.01), by species (p < 0.001), and by the interaction of both factors (p < 0.001) (Table 2). Cultivation in the plain area resulted in higher essential oil yield when compared to the mountainous area (1.38 ± 0.04% and 1.17 ± 0.04%, respectively), when the species factor was not considered. When comparing all the species regardless of altitude, laurel had the highest (2.68 ± 0.33%) and pelargonium had the lowest (0.42 ± 0.10 %) EO yield. Similar results were observed when considering the combined effect of the tested factors, where laurel and pelargonium plants grown in the mountainous area resulted in the highest and lowest EO yield (3.30 ± 0.11% and 0.19 ± 0.04% for laurel and pelargonium, respectively), which indicated the significance of the genotype on this parameter, apart from the growing location. In particular, the high altitude of the mountainous area increased by 16.5% and by 60.6% for the EOs yield of artemisia and laurel, respectively, when compared to plants grown in the plain area. In contrast, EOs yield of pelargonium, spearmint, lavender, and lemon verbena decreased significantly (a decrease of 84.1%, 53.9%, 45.3%, and 30.7%, respectively) in plants grown in high altitudes when compared to plants grown in the plain area. Therefore, EOs yield was impacted by the species and altitude in a variable manner for most of the studied MAPs, whereas, in rosemary and sage, EO yield was not affected by altitude and averaged at 1.03% and at 1.89% (for rosemary and sage, respectively) (Table 2). Mahomoodally et al. [74] reported that EO yield can alter during the different months of the year and can be decreased in areas that receive less solar radiation, while Khorshidi et al. [75] reported that plant density and nutrient availability may also affect plant growth and EOs yield as in the case of mountainous plants of our study. Moreover, a different altitude is affecting not only the antioxidant but also the antimicrobial properties of EOs of Thymus capitata (L.) due to differences in volatile compounds’ profile and in contents of bioactive substances [76]. The lowest EO yield in Tussilago farfara (L.) reported at low altitudes (i.e., 229 m) also revealed the highest antioxidant activity when compared to the plants grown in higher ones [48]. These findings agree with the results of our study where a decreased EO yield and increased antioxidants content was found in laurel in the plain area, but not in pelargonium and lemon verbena. This contradiction could be attributed not only to the different altitude and the climatic conditions of each location but also to differences among the studied MAP species. According to Maurya et al. [77] and Bailen et al. [78], the metabolic pathways related to essential oils composition may be affected by both environmental and genetic factors. Moreover, Formisano et al. [79] reported that chamomile harvested at low altitudes (i.e., 81–89 m) revealed increased EOs yield when compared to plants harvested from higher altitudes (i.e., 640–675 m), which is in accordance with our findings for pelargonium, spearmint, lavender, and lemon verbena (Table 2).
The effect of altitude on the EOs chemical composition of the examined MAP species is given in Table 3, Table 4, Table 5, Table 6, Table 7, Table 8, Table 9, Table 10 and Table 11. In the case of artemisia, EOs’ analysis revealed the presence of 28 and 30 individual compounds, which represent a total percentage of ≥92.01% of the oil profile for the plain and mountainous plants, respectively. The most abundant class (42.69% and 47.75%) was oxygenated monoterpenes, which was followed by oxygenated sesquiterpenes (39.03% and 31.01%), monoterpenes hydrocarbon (10.70% and 11.77%), and sesquiterpenes hydrocarbons (1.35% and 0.98%) for the plain and mountainous plants, respectively (Table 3). The major constituents of the examined artemisia EOs in decreasing order were 1,8-cineole (19.63–27.02%), cis-dihydroagarofuran (11.74–13.00%), silphiperfol-5-en-3-one A (9.71–13.06%), borneol (10.88–11.08%), camphor (5.92–8.59%), and p-cymene (5.69–7.26%). Camphene, ascaridole, silphiperfol-5-en-3-one B, silphiperfol-5-en-3-ol A, silphiperfol-5-en-3-ol B, presilphiperfol-8-ol, and caryophylla-4(12),8(13)-dien-5b-ol varied between 1–4%, while the rest of the compounds were identified in amounts lower than 1% of the total volatile components content (Table 3). Artemisia plants grown in the mountain had significantly higher content of 1,8-cineole and p-cymene but lower camphor and camphene content when compared to the plants grown in the plain (Table 3). The essential oil composition of artemisia has been widely studied and the literature reports show a great variability in essential oil composition where the main detected compounds were trans-sabinyl acetate and α-terpineol [80], davanone derivatives, and 4-Methyl-pent-2-enolid [81], whereas Mucciarelli et al. [82] reported 1,8-cineole as the main compound in the case of our study.
In laurel, EOs analysis resulted in 32 individual compounds in total for both the plain and mountainous plants, which represent ≥99.34% of the total oil profile (Table 4). Oxygenated monoterpenes were the most abundant class (78.08% and 64.70%), which is followed by monoterpenes hydrocarbon (11.91% and 20.04%), and by oxygenated sesquiterpenes (0.41% and 0.13%), as sesquiterpenes hydrocarbons were not detected in either plain or mountainous plants (Table 4). The main laurel component was 1,8-cineole (56.63–69.48%), which was followed by the α-terpinyl acetate (7.19–13.07%) while α-pinene, sabinene, β-pinene, p-cymene, terpinen-4-ol, myrtenal, carvone, and δ-terpinyl acetate that varied between 1–4% (Table 4). Laurel plants grown in the mountain had significantly higher α-pinene, sabinene, β-pinene, and α-terpinyl acetate but lower 1,8-cineole and p-cymene compared to the relevant plants grown in the plain area (Table 4). 1,8-cineole and α-terpinyl acetate was also reported as the major volatile compound in laurel essential oils by Ordoudi et al. [83] and Fidan et al. [84], while other studies refer to 1,8-cineole and linalool [85], eucalyptol, and terpinyl acetate [86], since extraction methods and sample preparation may affect the essential oils’ composition [87,88].
The EOs’ composition of lavender aerial parts is presented in Table 5 with 35 and 32 compounds identified in plain and mountainous plants, respectively, which represent ≥99.49% of the total oil profile. The classification of individual components revealed the oxygenated monoterpenes (87.39% and 89.28%) as most abundant, which is followed by monoterpenes hydrocarbon (5.26% and 5.45%), oxygenated sesquiterpenes (3.10% and 1.88%), and, lastly, by sesquiterpenes hydrocarbons (0.76% and 0.33%) for the plain and mountainous plants, respectively (Table 5). The main constituents of lavender EOs in decreasing order were 1,8-cineole (30.82–45.31%), which is followed by camphor (30.48–34.29%), linalool (0.30–7.47%), borneol (5.48–6.34%), and carvone (1.34–5.53%), whereas α-pinene, camphene, β-pinene, p-cymene, limonene, terpene-4-ol, caryophyllene oxide, and tau-cadinol levels varied between 1–4% (Table 5). In mountainous conditions, lavender EOs had higher camphor, linalool, and terpene-4-ol, but lower 1,8-cineole and p-cymene content compared to plants harvested in the plain area (Table 5). Contrasting reports exist in the literature regarding the essential oil composition of lavender, where linalyl acetate and linalool [89,90] or terpinene-4-ol [91] were identified as the major compounds, since agronomic practices may affect essential oil composition [89]. Moreover, Łyczko et al. [92] reported that camphor is an important quality marker for lavender essential oil. The higher the ratio of linalool and linalyl acetate is to camphor, the better the quality is. They also suggested that drying methods may significantly affect the essential oil composition. Similarly to our study, Oroian et al. [93] reported that geographical origin of lavender samples has a significant effect on essential oils composition, even though they reported linalool and linalyl acetate as the major compounds.
In lemon verbena, EOs’ analysis resulted in 30 and 31 individual compounds for the plain and mountain grown plants, respectively, which represented ≥98.99% of the total oil profile (Table 6). The analysed oils were dominated by the monoterpenes fraction. In particular, the oxygenated monoterpenes were the most represented class with percentages of 49.72% and 69.45%, which were followed by monoterpenes hydrocarbon (19.73% and 14.60%), oxygenated sesquiterpenes (17.50% and 8.13%), and sesquiterpenes hydrocarbons (10.87% and 6.28%) for the plain and mountainous plants, respectively. The main constituents for lemon verbena were geranial (α trans citral) (22.42–29.06%), neral (β cis citral) (16.03–17.72%), limonene (8.81–15.67%), caryophyllene oxide (4.67–9.38%), 1,8-cineole (6.85–8.41%), ar-curcumene (5.46–7.42%), spathulenol (3.08–6.45%), camphor (1.35–5.87%), α-thujone (1.13–5.25%), while the α-pinene, sabinene, β-thujone, and β-caryophyllene varied between 1–4% (Table 6). Lemon verbena grown in the mountain had significantly lower limonene, sabinene, β-caryophyllene, spathulenol, and caryophyllene oxide when compared to the relevant plants grown in the plain area (Table 6). The same compounds have been identified as the major constituents of lemon verbena EOs in various reports [94]. However, a seasonal effect on EOs was also suggested [95,96].
Pelargonium EOs’ analysis is presented in Table 7 and it revealed the identification of 29 and 27 components in plain and mountainous plants, respectively, that represented ≥99.65% of the total oil profile. Following constituents’ classification, the oxygenated monoterpenes predominated (76.20% and 73.04%) and was followed by oxygenated sesquiterpenes (10.19% and 8.13%), monoterpenes hydrocarbon (0.23% and 7.97%), and sesquiterpenes hydrocarbons (4.01% and 0.75%) for the plain and mountainous plants, respectively. The most abundant oil component was citronellol (24.25–36.69%), which was followed by geraniol (11.13–15.45%), citronellyl formate (13.29–14.11%), isomenthone (5.78–10.61%), geranyl formate (4.11–4.75%), γ-eudesmol (5.92–6.92%), cis-rose oxide (1.89–5.79%), and α-pinene (0.23–6.64%). The oil components of artemisia ketone, linalool, trans-rose oxide, germacrene D, phenethyl tiglate, geranyl tiglate, and farnesyl acetone varied from 1–4%, while other compounds were identified in amounts lower than 1% of the total volatile components content (Table 7). Compared to the plain area, mountainous plants contained significantly higher levels of α-pinene, artemisia ketone, cis-rose oxide, trans-rose oxide, isomenthone, phenethyl tiglate, and lower levels of citronellol, geraniol, γ-eudesmol, and geranyl tiglate (Table 7). Similarly to our study, citronellol and geraniol were identified as the major compounds of pelargonium EOs in many other reports, which also correlate with the bioactive properties of the EOs of the species with the presence of these compounds [97,98,99,100].
In rosemary, EOs analysis revealed the presence of 29 individual compounds for both plain and mountainous plants, which represent ≥99.87% of the total oil profile (Table 8). The main detected class was that of oxygenated monoterpenes (67.48% and 64.33%), which was followed by monoterpenes hydrocarbon (30.97% and 32.96%) for the plain and mountainous plants, respectively, whereas very low amounts of sesquiterpenes (oxygenated up to 0.13% and hydrocarbons up to 1.36%) were identified (Table 8). Accordingly, the major oil constituents in decreasing order were 1,8-cineole (32.94%), camphor (19.21–20.86%), α-pinene (12.01–13.05%), camphene (8.12–8.29%), and borneol (7.88–8.94%), whereas β-pinene, β-myrcene, p-cymene, limonene, and α-terpineol varied between 1–4% (Table 8). Rosemary plants grown in the mountain had significantly lower camphor, and α-terpineol content compared to the plants grown in the plain. The detected profile of volatile compounds is typical for the species based on several literature reports [101,102], whereas Contini et al. [103] reported a-pinene and 1,8-cineole as the most abundant compounds. Moreover, according to Sabbahi et al. [104], the profile of the major compounds was not affected by the elevation gradient. Compositional variability is attributed mostly to genetic factors.
Sage EOs’ analysis revealed the identification of 31 and 32 individual components for the plain and mountainous plants, respectively, which represent ≥99.97% of the total oil profile (Table 9). The most abundant class was oxygenated monoterpenes (73.20% and 65.53%), which was followed by monoterpenes hydrocarbon (20.35% and 22.58%), sesquiterpenes hydrocarbons (1.36% and 5.38%), and oxygenated sesquiterpenes (3.55% and 2.23%) for the plain and mountain plants, respectively (Table 9). The main oil constituents in decreasing order were camphor (22.26% and 16.98%), 1,8-cineole (16.69% and 15.28%), α-thujone (23.83% and 5.34%), β-thujone (5.23% and 13.32%), borneol (3.54% and 13.01%), camphene (7.10% and 8.13%), and α-pinene (4.33% and 5.62%) (Table 9). Moreover, EOs of sage plants grown in the mountain had significantly higher content of α-pinene, limonene, β-thujone, borneol, and manool, but lower α-thujone, camphor, and viridiflorol content when compared to the plants grown in the plain area. According to Bedini et al. [105], a-thujone was the major compound of sage EOs, while they also detected significant amounts of camphor and 1,8-cineole. Moreover, EOs’ composition may be affected by several factors such as environmental stressors [106], or the application of biofertilizers and bio-stimulants [107], which indicated the importance of exogenous factors on EOs’ biosynthesis. In contrast, Cvetkovikj et al. [108] who studied several sage populations from Balkan countries identified four distinct chemotypes, which differ in cis-thujone, trans-thujone, and camphor content and suggested a significant correlation of essential oil composition with geographic variables.
Spearmint EOs analysis resulted in the presence of 33 and 34 individual compounds for the plain and mountainous plants, respectively, which represented ≥99.91% of the total oil profile (Table 10). The most abundant class was oxygenated monoterpenes (79.59% and 74.03%), which were followed by monoterpenes hydrocarbon (15.95% and 10.58%), sesquiterpenes hydrocarbons (3.01% and 2.81%), and very low amounts of oxygenated sesquiterpenes for the plain and mountainous plants, respectively (Table 10). The major constituents of the examined spearmint EO in decreasing order were carvone (72.12% and 50.12%), limonene (12.07% and 6.65%), 1,8-cineole (4.98% and 5.68%), cis-dihydro carvone (0.65% and 12.97%), dihydrocarveol acetate (0.43% and 5.67%), and cis-carvyl acetate (0.27% and 4.92%). Spearmint plants grown in the mountain had significantly higher content of 1,8-cineole, cis-dihydro carvone, neo-dihydro carveol, cis-carveol, dihydrocarveol acetate, and cis-carvyl acetate but lower carvone, and limonene when compared to the plants grown in a plain (Table 10). Carvone was the most abundant spearmint EO constituent in several other reports [109,110], while agronomic factors such as salinity and water stress may affect EOs yield and composition [64,111].

3.3. Correlation of Antioxidant and Reducing Activity with Polyphenols and Essential Oils Components

MAP species are highly appreciated as a natural source of antioxidants, while phenolic compounds and essential oils components are involved in such antioxidant capacity [34,46]. To evaluate the contribution of phenols, flavonoids, flavanols, and essential oils components, only the five most abundant constituents of each EOs were considered. For their total antioxidant capacity, linear correlation coefficients were determined and presented in detail in Tables S2–S9. The correlation coefficient (r) and p-values between the analysed EO compounds and the antioxidant capacity of artemisia are given in Table S2. In the plain area, phenols content was strongly and positively correlated with flavanols. DPPH was correlated with borneol, while flavonoids were correlated with DPPH and cis-dihydroagarofuran. Accordingly, for plants grown in the mountainous area, a positive correlation was found with phenols and antioxidant activity of DPPH and FRAP. Borneol has been associated with antioxidant activity as well as with antihypertensive properties in animal models [112,113]. Its increased content could increase the overall antioxidant capacity of artemisia. Moreover, in the study of Wang et al. [114], it was observed that thyme borneol essential oils presented the highest antioxidant activities (reducing power and β-carotene bleaching) and the highest total phenols content among 26 essential oils.
In pelargonium in the plain area, phenols were positively correlated with DPPH, flavonoids, and flavanols. FRAP was correlated to flavonoids and flavanols, while flavonoids were correlated to flavanols (Table S3). In the mountainous area, phenols were correlated to DPPH, FRAP, flavonoids, flavanols, and EO yield. DPPH was correlated to FRAP and flavanols while FRAP was correlated to flavonoids and flavanols. Essential oil yield was correlated to the geraniol content (Table S3), as decreased EO and geraniol content were observed in the mountainous areas.
In laurel grown in plain areas, phenols were positively correlated to DPPH, FRAP, flavonoids, and flavanols. DPPH was positively correlated with flavonoids and flavanols and negatively correlated with 1,8-cineole. FRAP was positively correlated with flavonoids, flavanols, and β-pinene and negatively correlated with 1,8-cineole. Flavonoids were correlated with flavanols, while flavanols were positively correlated with α-pinene (Table S4). In the mountainous area, phenols were positively correlated with DPPH and FRAP. FRAP was positively correlated to flavonoids and flavanols. Flavonoids were correlated with flavanols and 1,8-cineole. It has been reported that lavender plants subjected to salinity stress in hydroponics showed increased levels of α-pinene (4.45% and 3.86% 100 mM NaCl) in conjunction with the increased levels of antioxidants [(DPPH, FRAP, ABTS (2,2’-azino-bis-3-ethylbenzthiazoline-6-sulphonic acid), and phenols)] [115]. Additionally, Sideritis perfoliata plants that were cultivated under organic cultivation system appeared to have higher antioxidant activity in terms of DPPH, ABTS, FRAP, and total phenolic content and higher content in α-pinene and β-pinene (25.35% and 27.98% for α-pinene and 6.51% and 7.13% for β-pinene) [46,116]. According to Salehi et al. [117], pinenes possess significant bioactive properties and the increased content in mountainous grown plant of laurel observed could partially justify the increased antioxidant activity. Moreover, the low antioxidant activity of 1,8-cineole [117,118] may justify the negative correlation with FRAP and DPPH values observed in our study.
Correlation analysis in lavender revealed that, in plain grown plants, phenols were positively correlated with DPPH, FRAP, and flavonoids. DPPH was positively correlated with flavonoids and FRAP was positively correlated with flavonoids. Flavonoids were positively correlated with 1,8-cineole. Flavanols were positively correlated with borneol (Table S5). Ιn the case of mountainous grown lavender, phenols were positively correlated with DPPH, FRAP, and flavonoids. DPPH was positively correlated with FRAP and FRAP was positively correlated with flavonoids. However, flavonoids were negatively correlated with carvone. Carvone isolated from spearmint exhibited diverse biocidal activities including antioxidant, insecticidal, antifungal, and antibacterial as reviewed by Elmastaş et al. [109].
In lemon verbena grown in plain areas, phenols were positively correlated to DPPH, and flavonoids. DPPH was positively correlated with FRAP and flavonoids. FRAP was positively correlated to flavonoids. Flavonoids were positively correlated with neral (Table S6). In the mountainous area, lemon verbena’s phenols were positively correlated with FRAP. FRAP was positively correlated with flavonoids, 1,8-cineole, and negatively correlated with D-limonene, neral, geranial, and caryophyllene oxide. 1,8-cineole exhibited insecticidal and antimicrobial, anti-allergic and anti-inflammatory, hepatoprotective, antitumoral, and gastroprotective action as reviewed by Caldas et al. [119]. In contrast to our study, D-limonene has been associated with significant antioxidant activities [120,121], which is a finding that could be associated with possible interactions among the essential oil components as already reported for rosemary essential oils by Wang et al. [122]. Similar assumptions could be made for geranial, neral, and caryophyllene oxide [123,124].
In rosemary grown in plain areas, phenols were positively correlated with FRAP, flavonoids, and camphor and negatively correlated with α-pinene. FRAP was positively correlated with flavonoids, while flavonoids were negatively correlated with 1,8-cineole (Table S7). In the mountainous area, rosemary phenols were positively correlated with DPPH, FRAP, and flavonoids, but negatively correlated with borneol. DPPH was positively correlated with FRAP and flavonoids. FRAP was positively correlated with flavonoids, while flavonoids were negatively correlated with borneol. The essential oil yield was negatively correlated with 1,8-cineole.
In sage grown in plain areas, phenols were positively correlated with FRAP and flavonoids. DPPH was negatively correlated with α-thujone. FRAP was positively correlated with flavonoids (Table S8). In the mountainous area, sage phenols were positively correlated with FRAP and flavonoids. FRAP was positively correlated with flavonoids. Flavanols were positively correlated with 1,8-cineole. EOs were positively correlated with camphor. When sage subjected to drought stress, total phenols, DPPH, FRAP, ABTS, flavonoids, and EO yield increased and this was reflected to the increased levels of camphor (45.91% full irrigation and 47.93% water deficit) and a-thujone (5.40% water deficit and 1.90% full irrigation) when compared with the plants subjected to full irrigation [125]. Similarly, increased camphor levels were found in Artemisia herba-alba plants as well as increased levels of total phenols and antioxidants [126]. It is widely known that high levels of camphor are toxic, whereas natural borneol is non-toxic [127]. 1,8-cineole also appeared to increase (from 18.61% to 35.42%) along with the increase in salinity levels, while the antioxidant activity and total phenol content of the methanolic extracts of Salvia mirzayanii plants also increased in saline conditions [128]. α-thujone exhibited antibacterial, cytotoxic, and antiviral activities [129]. Regarding the negative correlation of DPPH values with α-thujone, it has been previously reported that the specific compounds show low antioxidant activity [130], while it is also considered harmful to human health [131].
In the case of spearmint, correlation analysis in plain revealed that phenols were positively correlated with DPPH and FRAP, while DPPH was positively correlated with FRAP (Table S9). In the mountainous area, phenols were positively correlated with FRAP and flavonoids, while DPPH and FRAP were positively correlated with flavonoids. Flavonoids were positively correlated with flavanols, while flavanols were positively correlated with 1,8-cineole (Table S9). Spearmint plants exposed to multiple stress of salinity and copper toxicity revealed oxidative damage and decreased the levels of antioxidants and the levels of 1,8-cineole (4.46% and 3.55% sal and Cu) in leaves when compared with plants grown without stress [132].
The most important correlations related to antioxidant activity and chemical composition and essential oil compounds content are summarized in Table 11. The correlation analysis of antioxidant activity (DPPH and FRAP) with the major essential oil components and phenolic compounds (total phenols, flavonoids, and flavanols) showed a strong positive correlation of FRAP and DPPH values with total phenols, flavonoids, and flavanols content. However, this correlation varied depending on the species and the growing conditions (plain or mountainous area), which was also reflected in the results presented in Table 2. Moreover, in limited cases, antioxidant activity (DPPH or FRAP values) was positively correlated with essential oil components, as in the case of borneol and β-pinene in artemisia and laurel plants grown in the plain, respectively, or 1,8-cineole in verbena plants grown in the mountainous area. Similarly, antioxidant activity was negatively correlated with 1,8-cineole and α-thujone in laurel and sage plants grown in the plain as well as with neral, geranial, and caryophyllene oxide in lemon verbena plants grown in the mountainous area. These findings indicate that essential oils may contribute to the overall antioxidant activity depending on the species and its growing environment. Correlation analysis in our study was performed only with data from the most abundant compounds of the essential oils of each species, which eliminates the effect of minor compounds in terms of their antioxidant capacity. It is well known that, in natural matrices, synergistic effects may exist among their components and minor compounds can become essential for the antioxidant capacity of plant tissues [40,133,134,135,136]. Therefore, essential oils may exhibit higher antioxidant activity than the isolated components, as already reported by Wang et al. [117] for rosemary essential oils. In addition, essential oils may contain conjugated double bonds and phenolic groups with associated functional and antioxidant properties [135].

4. Conclusions

In the present study, we examined the levels of total phenolics, flavonoids, and flavanols content and antioxidant activity of eight MAP species, as affected by the environmental condition-altitude (mountainous versus plain areas) and their interconnection with the essential oil yield and composition. Altitude increased the phenolic compounds content and the antioxidant capacity of specific species such as artemisia, spearmint (total phenols only), and rosemary (flavonoids only), but higher antioxidant capacity was observed in plants grown in the plain compared to the mountainous area. EO yield was also affected by high altitude, by causing increased EO yield in laurel and decreased EO yield in pelargonium, and spearmint in mountainous areas. EO composition was also altered by the altitude. Plant antioxidant activity was positively correlated with total phenols, flavonoids, and flavanols and, in some cases, with specific constituents of the species (i.e., 1,8-cineole in laurel, α-thujone in sage, etc.), even though a varied response was observed depending on the species and the altitude. In conclusion, the effect of growing conditions defined by different environmental conditions-altitudes may significantly affect antioxidant compounds content and EO yield as well as composition of the studied MAPs in a species-dependent manner. These findings can be used to identify specific locations and ecosystems in which cultivation of MAPs could be introduced for producing high added value products with improved quality and bioactive properties.

Supplementary Materials

The following are available online at Tables S1–S9.

Author Contributions

Conceptualization, A.C. and N.T. Methodology, M.M. and A.C. Software, A.C. and M.M. Validation, M.M., A.C., and N.T. Formal analysis, A.C. Investigation, A.C. and M.M. Resources, N.T. Data curation, A.C., S.P., and N.T. Writing—original draft preparation, A.C., S.P., and N.T. Writing—review and editing, A.C., S.P., and N.T. Visualization, N.T. Supervision, N.T. Project administration, N.T. Funding acquisition, N.T. All authors have read and agreed to the published version of the manuscript.


This research has been co-financed by the start-up grant for the research project SALTAROMA in Cyprus University of Technology, by the project AgroLabs that has been developed under the Programme Interreg V-B Balkan - Mediterranean 2014–2020, co-funded by the European Union and National Funds of the participating countries. Cyprus University of Technology Open Access Author Fund.


We thank the Cypriot National Agricultural Department for providing the plant material at the plain area

Conflicts of Interest

The authors declare no conflict of interest.


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Table 1. Plant species and material used.
Table 1. Plant species and material used.
Common NameLatin NameFamilyPlant MaterialReported Medicinal
Artemisia Artemisia abrotanum LAsteraceaeLeavesAntifungal, anticancer, antiviral antibacterial, antioxidant, anaemia, amenorrhoea, anorexia, chronic fever, hepatitis, splenitis, hysteria [51].
Pelargonium Pelargonium roseum L.GeraniaceaeLeavesAntibacterial, antifungal, antioxidant, antitumor, nematocidal, intestinal problems, wounds and respiratory ailments, help hormonal balance, discharge toxins from liver, digestive [52].
nobilis L.
LauraceaeLeavesAntibacterial, antifungal, cytotoxicity, antioxidant, diuretic, gastrointestinal problems, to treat epilepsy, neuralgia, and parkinsonism [53].
Rosemary Rosmarinus officinalis L.LamiaceaeStem/
Antibacterial, antihepatotoxic, anti-tumour, anti-inflammatory, anti-trypanosomal, antispasmodic, immune stimulant activity, rheumatic complaints and circulatory disorders, tiredness, defective memory, carminative, rubefacient, promote digestion [54].
Spearmint Mentha spicata L.LamiaceaeStem/
Anti-inflammatory, sedative, antimicrobial, antioxidant, carminative, antispasmodic, diuretic, insecticidal [55].
Lavender Lavandula angustifolia L.LamiaceaeStem/
Antibacterial, insecticidal, sedative, analgesic, cytotoxic, anxiolytic, alleviate depression, headaches, and anxiety [56].
Lemon verbena Aloysia triphylla L.VerbenaceaeStem/
Antibacterial, antifungal, antioxidant, treatment of colic, diarrhea, indigestion, insomnia, anxiety, asthma, fever [57].
Sage Salvia officinalis L.LamiaceaeStem/
Antibacterial, antifungal, anticancer, antiviral, antidiabetic, antimutagenic, antiprotozoal, antidementia, antioxidant, anti-inflammatory [58].
Jacovides, C.P.; Timvios, F.S.; Papaioannou, G.; Asimakopoulos, D.N.; Theofilou, C.M. Ratio of PAR to broadband solar radiation measured in Cyprus. Agric. For. Meteorol. 2004, 121, 135–140.
Table 2. Effects of altitude (mountain vs. plain) on the content of total phenols (μmol GAE g−1 fw), total flavonoids (mg rutin g−1 fw), total flavanols (mg catechin g−1 fw), antioxidant and reducing activity (DPPH, FRAP, mg trolox g−1 fw), and essential oil (EO) yield (%) in selected medicinal plant species.
Table 2. Effects of altitude (mountain vs. plain) on the content of total phenols (μmol GAE g−1 fw), total flavonoids (mg rutin g−1 fw), total flavanols (mg catechin g−1 fw), antioxidant and reducing activity (DPPH, FRAP, mg trolox g−1 fw), and essential oil (EO) yield (%) in selected medicinal plant species.
SpeciesAltitudeTotal PhenolsFlavonoidsFlavanolsDPPHFRAPEO
Plain 78.42 ± 6.62A7.01 ± 0.78A7.16 ± 2.29A25.02 ± 3.72A12.31 ± 1.24A1.38 ± 0.04A
Mountain68.68 ± 4.23A7.90 ± 1.14A2.49 ± 0.63B13.53 ± 0.63B11.53 ± 0.84A1.17 ± 0.04B
Total mean73.55 ± 3.947.46 ± 0.694.82 ± 1.2019.27 ± 1.9711.92 ± 0.751.28 ± 0.03
Artemisia 73.16 ± 12.81BC13.52 ± 2.71B0.44 ± 0.13C12.64 ± 1.63C9.57 ± 1.48C0.90 ± 0.08CD
Pelargonium 112.03 ± 17.69A3.65 ± 0.73D29.14 ± 5.78A47.70 ± 10.41A25.28 ± 1.91A0.42 ± 0.10D
Laurel 113.02 ± 5.31A8.15 ± 1.25C8.48 ± 0.74B32.51 ± 5.53B15.59 ± 1.22B2.68 ± 0.33A
Rosemary 83.89 ± 3.84B4.66 ± 0.99CD0.28 ± 0.16C15.34 ± 1.38C12.95 ± 0.87B1.03 ± 0.05C
Spearmint 52.28 ± 4.50CD8.18 ± 1.13C0.00 ± 0.00C11.27 ± 1.02C6.84 ± 1.22CD1.89 ± 0.09B
Lavender 61.06 ± 2.77BC17.31 ± 1.03A0.02 ± 0.00C16.06 ± 0.69C13.24 ± 0.80B0.63 ± 0.02CD
Lemon verbena 33.47 ± 1.37D2.24 ± 0.27D0.22 ± 0.05C6.13 ± 0.32C3.66 ± 0.23D0.76 ± 0.11CD
Sage 59.51 ± 4.37BC2.06 ± 0.68D0.04 ± 0.01C12.54 ± 1.26C8.23 ± 0.89C1.90 ± 0.31B
Total mean 73.55 ± 3.947.46 ± 0.694.82 ± 1.2019.27 ± 1.9711.92 ± 0.751.28 ± 0.12
Artemisia Plain 40.6 ± 2.4ijkY4.7 ± 0.8efghi0.06 ± 0.02d7.5 ± 1.0fg4.8 ± 0.6fg0.58 ± 0.02g
Mountain105.7 ± 17.1c22.3 ± 1.0a0.8 ± 0.1d17.8 ± 0.3c14.3 ± 0.5c0.68 ± 0.01efg
Pelargonium Plain 166.9 ± 12.0a5.9 ± 0.6defg47.1 ± 3.8a80.9 ± 6.0a29.9 ± 2.1a0.65 ± 0.07fg
Mountain57.1 ± 5.2ghij1.4 ± 0.2ij11.1 ± 1.7b14.5 ± 1.0cde20.6 ± 1.7b0.19 ± 0.04e
Laurel Plain 126.0 ± 6.6b11.3 ± 1.5c9.3 ± 1.2bc49.6 ± 4.1b18.9 ± 1.0b2.05 ± 0.39c
Mountain100.0 ± 3.5cd5.0 ± 0.8efgh7.7 ± 0.8c15.4 ± 0.8cde12.3 ± 1.1cd3.30 ± 0.11a
Rosemary Plain 87.0 ± 4.7cde2.7 ± 1.2ghij0.56 ± 0.02d14.4 ± 2.7cde12.8 ± 1.5cd1.08 ± 0.07de
Mountain80.8 ± 6.2def6.6 ± 1.2defnd16.3 ± 1.0cd13.1 ± 1.0cd0.99 ± 0.08def
Spearmint Plain 41.9 ± 3.5ijk7.4 ± 1.8dend9.0 ± 0.9efg5.2 ± 0.7fg2.61 ± 0.04b
Mountain62.7 ± 5.8fgh8.8 ± 1.5cdnd13.5 ± 1.3cdef8.5 ± 2.2ef1.20 ± 0.06d
Lavender Plain 58.1 ± 3.3ghi17.0 ±1.0b0.90 ± 0.05d16.4 ± 0.8cd12.6 ± 0.9cd0.99 ± 0.13def
Mountain64.0 ± 4.4fgh17.2 ± 1.9b0.91 ± 0.05d15.7 ± 1.2cde13.9 ± 1.4cd0.54 ± 0.01ge
Lemon verbena Plain 36.6 ± 1.8jk2.9 ± 1.6ghij0.15 ± 0.05d6.7 ± 0.4g4.1 ± 0.3g1.06 ± 0.07de
Mountain30.3 ± 1.1k1.6 ± 0.3hij0.28 ± 0.09d5.5 ± 0.4g3.2 ± 0.2g0.73 ± 0.07efg
Sage Plain 70.1 ± 4.9efg3.8 ± 0.8fghi0.08 ± 0.04d15.7 ± 1.5cde10.14 ± 1.05de2.06 ± 0.02c
Mountain48.9 ± 3.9hijk0.29 ± 0.03jnd9.4 ± 0.9defg6.32 ± 0.97fg1.72 ± 0.13c
InteractionS x A******************
Y values (means ± SE, n = 6) in columns corresponding to the main factors (Altitude and Species) followed by the same uppercase letter, and values corresponding to the interaction of the main factors (Altitude and Species), which is followed by the same lowercase letter, are not significantly different, p ≤ 0.05. nd: not detected. ns, **, and *** indicate non-significant or significant differences at p < 0.01, and p < 0.001, respectively, following a two-way ANOVA.
Table 3. Chemical composition (%) of essential oils of artemisia plants grown at different altitudes (plain vs. mountain). Data are expressed as means ± SE (n = 3), and significant differences (p < 0.05, p < 0.01, p < 0.001) in bold between treatments are indicated by Student’s t-test p-values.
Table 3. Chemical composition (%) of essential oils of artemisia plants grown at different altitudes (plain vs. mountain). Data are expressed as means ± SE (n = 3), and significant differences (p < 0.05, p < 0.01, p < 0.001) in bold between treatments are indicated by Student’s t-test p-values.
ComponentsRIPlainMountainStudent’s t-Test
α-Pinene9330.19 ± 0.0970.39 ± 0.1590.343
Camphene9483.63 ± 0.1512.76 ± 0.026<0.05
β-Pinene9770.33 ± 0.0170.60 ± 0.2480.339
α-Terpinene10170.86 ± 0.1590.26 ± 0.015<0.05
p-Cymene10245.69 ± 0.4667.26 ± 0.364<0.05
β-Phellandrene10290.00± 0.0000.12 ± 0.0720.163
1,8-Cineole103119.63 ± 1.40027.02 ± 0.341<0.01
γ-Terpinene1058nd0.37 ± 0.003-
cis-Sabinene hydrate10670.23 ± 0.123nd0.132
trans-Sabinene hydrate11000.27 ± 0.144nd0.131
cis-p Menth-2-en-1-ol 1121nd0.41 ± 0.017-
trans-p Menth-2-en-1-ol 11380.44 ± 0.020.54 ± 0.017<0.05
Camphor11458.59 ± 0.6075.92 ± 0.12<0.05
Borneol116611.08 ± 0.6810.88 ± 0.070.789
Terpinen-4-ol11780.74 ± 0.0261.11 ± 0.026<0.001
Ascaridole12381.7 ± 0.5360.82 ± 0.0090.174
cis-Piperotone epoxide 1254nd0.31 ± 0.012-
trans-Piperotone epoxide 1257nd0.57 ± 0.009-
Isobornyl acetate12850.59 ± 0.1370.49 ± 0.0200.537
Carvacrol1300nd0.16 ± 0.095-
Silphiperfol-5-ene13240.46 ± 0.0700.61 ± 0.0460.155
Presilphiperfol-7-ene1336nd0.08 ± 0.043-
7-epi Silphiperfol-5-ene 13430.21 ± 0.1110.07 ± 0.0400.312
Silphiperfol-4.7(14)-diene13590.32 ± 0.0350.23 ± 0.0290.108
Germacrene D14970.36 ± 0.180nd-
Silphiperfolan-6a-ol15180.67 ± 0.0500.79 ± 0.0380.143
cis-Dihydroagarofuran 153313.00 ± 0.76711.74 ± 0.1910.187
Silphiperfol-5-en-3-ol B15441.90 ± 0.1281.56 ± 0.0320.058
Silphiperfol-5-en-3-one B15562.54 ± 0.0741.85 ± 0.020<0.001
Silphiperfol-5-en-3-ol A15622.02 ± 0.1561.55 ± 0.061<0.05
Silphiperfol-5-en-3-one A158113.06 ± 1.3209.71 ± 0.2940.068
Spathulenol15820.21 ± 0.207nd-
Presilphiperfol-8-ol15853.70 ± 0.1993.17 ± 0.0120.056
Caryophylla-4(12),8(13)-dien-5b-ol16381.22 ± 0.1520.65 ± 0.058<0.05
epi-α-Bisabolol 16850.70 ± 0.055nd-
Total Identified 94.35 ± 0.98192.01 ± 0.0440.075
Monoterpenes hydrocarbons 10.70 ± 0.42211.77 ± 0.1090.070
Sesquiterpenes hydrocarbons 1.35 ± 0.3210.98 ± 0.0080.319
Oxygenated monoterpenes 42.69 ±1.30147.75 ± 0.285<0.05
Oxygenated sesquiterpenes 39.03 ± 1.74031.01 ± 0.326<0.05
Others 0.58 ±0.1360.49 ± 0.0200.537
nd: not detected.
Table 4. Chemical composition (%) of essential oils of laurel plants grown at different altitudes (plain vs. mountain). Data are expressed as means ± SE (n = 3), and significant differences (p < 0.05, p < 0.01, p < 0.001) in bold between treatments are indicated by Student’s t-test p-values.
Table 4. Chemical composition (%) of essential oils of laurel plants grown at different altitudes (plain vs. mountain). Data are expressed as means ± SE (n = 3), and significant differences (p < 0.05, p < 0.01, p < 0.001) in bold between treatments are indicated by Student’s t-test p-values.
ComponentsRIPlainMountainStudent’s t-Test
α-Τhujene9260.07 ± 0.0430.10 ± 0.0500.709
α-Pinene9333.13 ± 0.1914.03 ± 0.102<0.05
Camphene9480.03 ± 0.0270.02 ± 0.0170.766
Sabinene9731.52 ± 0.1008.72 ± 0.067<0.001
β-Pinene9773.00 ± 0.1183.78 ± 0.044<0.01
Dehydro-1,8-cineole9910.56 ± 0.0400.52 ± 0.0700.647
α-Terpinene10170.14 ± 0.0340.34 ± 0.038<0.05
p-Cymene10242.97 ± 0.2520.70 ± 0.009<0.001
Limonene10280.69 ± 0.0671.44 ± 0.055<0.001
1,8-Cineole103169.48 ± 1.57756.63 ± 0.591<0.01
γ-Terpinene10580.36 ± 0.0620.77 ± 0.052<0.01
cis-Sabinene hydrate10670.06 ± 0.0380.37 ± 0.038<0.01
Terpinolene1089nd0.14 ± 0.015-
trans-Sabinene hydrate11000.10 ± 0.0520.50 ± 0.029<0.01
trans-p Mentha-2,8-dienol11190.15 ± 0.0770.18 ± 0.0960.800
trans-Pinocarveol11390.89 ± 0.0900.42 ± 0.096<0.05
Camphor11450.04 ± 0.0370.28 ± 0.032<0.01
Pinocarvone11630.80 ± 0.1200.30 ± 0.085<0.05
p-Mentha-1,5-dien-8-ol 11650.59 ± 0.1180.54 ± 0.0840.732
Terpinen-4-ol11782.78 ± 0.3641.98 ± 0.1870.123
Thuj-3-en-10-al11840.11 ± 0.0580.12 ± 0.0330.962
cis-Pinocarveol11860.43 ± 0.0600.07 ± 0.073<0.05
α-Terpineol11910.78 ± 0.1711.21 ± 0.320.299
Myrtenal11971.14 ± 0.1490.52 ± 0.115<0.05
trans-Carveol12190.11 ± 0.012nd-
cis-Carveol12310.47 ± 0.0580.13 ± 0.078<0.05
Carvone12440.49 ± 0.1071.21 ± 0.111<0.01
Bornyl acetate12850.03 ± 0.0330.02 ± 0.0170.678
δ-Terpinyl acetate 13161.02 ± 0.2220.95 ± 0.1390.802
α-Terpinyl acetate13497.19 ± 0.94813.07 ± 0.359<0.01
Eugenol1356nd0.14 ± 0.137-
Eugenol methyl14040.07 ± 0.073nd-
Caryophyllene oxide15870.11 ± 0.1070.03 ± 0.0270.507
β-Εudesmol 16510.31 ± 0.0430.10 ± 0.050<0.05
Total Identified 99.62 ± 0.18899.34 ± 0.1900.355
Monoterpenes hydrocarbons 11.91 ± 0.65920.04 ± 0.274<0.001
Sesquiterpenes hydrocarbons 0.00 ± 0.0000.00 ± 0.000-
Oxygenated monoterpenes 78.08 ±1.52864.70 ± 0.340<0.001
Oxygenated sesquiterpenes 0.41 ± 0.0840.13 ± 0.0680.059
Others 8.31 ±1.12614.04 ± 0.402<0.01
nd: not detected.
Table 5. Chemical composition (%) of essential oils of lavender plants grown at different altitudes (plain vs. mountain). Data are expressed as means ± SE (n = 3), and significant differences (p < 0.05, p < 0.01, p < 0.001) in bold between treatments are indicated by Student’s t-test p-values.
Table 5. Chemical composition (%) of essential oils of lavender plants grown at different altitudes (plain vs. mountain). Data are expressed as means ± SE (n = 3), and significant differences (p < 0.05, p < 0.01, p < 0.001) in bold between treatments are indicated by Student’s t-test p-values.
ComponentsRIPlainMountainStudent’s t-Test
α-Pinene9331.21 ± 0.3421.01 ± 0.0490.606
Camphene9481.23 ± 0.1831.12 ± 0.0320.575
Sabinene9730.22 ± 0.0150.07 ± 0.038<0.05
β-Pinene9771.07 ± 0.1931.15 ± 0.0530.698
p-Cymene10241.28 ± 0.0460.96 ± 0.031<0.01
Limonene10281.44 ± 0.3621.98 ± 0.1900.254
1,8-Cineole103145.31 ± 2.17730.82 ± 1.099<0.01
γ-Terpinene10580.04 ± 0.0370.17 ± 0.015<0.05
cis-Sabinene hydrate10670.31 ± 0.102nd-
Linalool11000.30 ± 0.1377.47 ± 2.038<0.05
α-Campholenal11270.06 ± 0.0630.08 ± 0.0420.805
trans-Pinocarveol11390.36 ± 0.1600.28 ± 0.0250.647
Camphor114530.48 ± 0.93534.29 ± 0.946<0.05
Pinocarvone11630.41 ± 0.1600.30 ± 0.0280.547
Borneol11665.48 ± 0.5206.34 ± 0.2390.206
Terpene-4-ol11780.48 ± 0.0521.36 ± 0.228<0.05
meta-p-Cymen-8-ol 11810.17 ± 0.091nd-
p-Cymen-8-ol11850.41 ± 0.0580.21 ± 0.029<0.05
Cryptone11870.96 ± 0.5130.90 ± 0.1120.905
α-Terpineol11910.68 ± 0.0830.84 ± 0.0830.244
Myrtenal11970.73 ± 0.3970.39 ± 0.0410.446
Verbenone12110.09 ± 0.087nd-
trans-Carveol12190.06 ± 0.0570.15 ± 0.0780.371
Bornyl formate1229nd0.25 ± 0.010-
Cumic aldehyde12410.75 ± 0.3770.93 ± 0.0500.655
Carvone12441.34 ± 0.0035.53 ± 1.591<0.05
Linalool acetate1255nd0.61 ± 0.296-
Bornyl acetate12850.21 ± 0.1070.07 ± 0.0700.326
Lavandulyl acetate12900.38 ± 0.0100.14 ± 0.070<0.05
α-Santalene14230.22 ± 0.1110.22 ± 0.0410.979
γ-Cadinene15250.55 ± 0.4230.11 ± 0.0560.364
Caryophyllene oxide15871.10 ± 0.2131.24 ± 0.0880.576
Cubenol16160.06 ± 0.060nd-
tau-Cadinol 16421.37 ± 0.7940.55 ± 0.0550.357
Bisabolol oxide II16560.24 ± 0.1190.04 ± 0.0430.201
α-Bisabolol16850.33 ± 0.1670.06 ± 0.0570.191
Muurol-5-en-4-one16890.21 ± 0.207nd-
Total Identified 99.49 ± 0.17799.67 ± 0.0680.405
Monoterpenes hydrocarbons 5.26 ± 0.1505.45 ± 0.2810.590
Sesquiterpenes hydrocarbons 0.76 ± 0.3200.33 ± 0.0280.249
Oxygenated monoterpenes 87.39 ±1.27989.28 ± 0.8900.293
Oxygenated sesquiterpenes 3.10 ± 0.7841.88 ± 0.1950.205
Others 1.76 ±0.4211.96 ± 0.3530.730
nd: not detected.
Table 6. Chemical composition (%) of essential oils of lemon verbena plants grown at different altitudes (plain vs. mountain). Data are expressed as means ± SE (n = 3), and significant differences (p < 0.05, p < 0.01, p < 0.001) in bold between treatments are indicated by Student’s t-test p-values.
Table 6. Chemical composition (%) of essential oils of lemon verbena plants grown at different altitudes (plain vs. mountain). Data are expressed as means ± SE (n = 3), and significant differences (p < 0.05, p < 0.01, p < 0.001) in bold between treatments are indicated by Student’s t-test p-values.
ComponentsRIPlainMountainStudent’s t-Test
α-Pinene9331.21 ± 0.1201.82 ± 0.6990.440
Camphene9480.42 ± 0.1182.00 ± 1.1550.244
Sabinene9731.87 ± 0.1190.68 ± 0.199<0.01
Oct-1-en-3-ol975nd0.19 ± 0.110-
β-Pinene9770.30 ± 0.0550.70 ± 0.4010.379
6-methyl-5 Hepten-2-one9840.36 ± 0.0450.06 ± 0.038<0.01
β-Myrcene9890.27 ± 0.0230.31 ± 0.1240.767
p Cymene1006nd0.28 ± 0.162-
Limonene102615.67 ± 0.6168.81 ± 1.091<0.01
1,8-Cineole10316.85 ± 0.3208.41 ± 2.3270.543
cis-Sabinene hydrate10670.30 ± 0.0060.09 ± 0.052<0.05
Linalool1100nd0.11 ± 0.066-
α-Thujone11061.13 ± 0.3125.25 ± 2.9210.233
β-Thujone11160.23 ± 0.1191.29 ± 0.7450.231
Camphor11451.35 ± 0.3415.87 ± 3.2300.236
Borneol11660.11 ± 0.1130.87 ± 0.3580.114
Isocitral11770.20 ± 0.1010.27 ± 0.1010.665
α-Terpineol11911.02 ± 0.0350.45 ± 0.139<0.05
Neral (β cis Citral)124016.03 ± 0.64217.72 ± 3.0140.612
Carvone12440.08 ± 0.0800.06 ± 0.0320.800
Geranial (α trans Citral)127122.42 ± 0.86629.06 ± 3.3980.131
α-Copaene13760.37 ± 0.025nd-
Geranyl acetate13830.81 ± 0.0290.82 ± 0.2280.967
β-Bourbonene14140.52 ± 0.0180.08 ± 0.046<0.001
α-Cedrene1422nd0.10 ± 0.055-
β-Caryophyllene14252.20 ± 0.2260.51 ± 0.188<0.01
Alloaromadendrene14640.37 ± 0.0300.13 ± 0.043<0.05
ar-Curcumene14967.42 ± 0.4515.46 ± 1.1030.176
Cubebol15270.28 ± 0.181nd-
Nerolidol E15680.32 ± 0.0060.16 ± 0.0920.159
Spathulenol15816.45 ± 0.2643.08 ± 0.921<0.05
Caryophyllene oxide15879.38 ± 0.7024.67 ± 0.990<0.05
Humulene epoxide II16080.28 ± 0.024nd-
epi-α-Cadinol16410.79 ± 0.0470.24 ± 0.136<0.05
Total Identified 98.99 ± 0.20699.53 ± 0.0890.075
Monoterpenes hydrocarbons 19.73 ± 0.83314.60 ± 1.125<0.05
Sesquiterpenes hydrocarbons 10.87 ± 0.7436.28 ± 1.431<0.05
Oxygenated monoterpenes 49.72 ±1.00669.45 ± 2.782<0.01
Oxygenated sesquiterpenes 17.50 ± 1.1168.13 ± 2.139<0.05
Others 1.16 ±0.0691.07 ± 0.3750.813
nd: not detected.
Table 7. Chemical composition (%) of essential oils of pelargonium plants grown at different altitudes (plain vs. mountain). Data are expressed as means ± SE (n = 3), and significant differences (p < 0.05, p < 0.01, p < 0.001) in bold between treatments are indicated by Student’s t-test p-values.
Table 7. Chemical composition (%) of essential oils of pelargonium plants grown at different altitudes (plain vs. mountain). Data are expressed as means ± SE (n = 3), and significant differences (p < 0.05, p < 0.01, p < 0.001) in bold between treatments are indicated by Student’s t-test p-values.
ComponentsRIPlainMountainStudent’s t-Test
α-Pinene9330.23 ± 0.0156.64 ± 0.372<0.001
β-Pinene977nd0.67 ± 0.015-
β-Myrcene989nd0.26 ± 0.006-
Limonene1028nd0.41 ± 0.009-
Artemisia ketone1059nd2.11 ± 0.041-
Linalool11001.02 ± 0.2250.88 ± 0.0180.578
cis-Rose oxide11101.89 ± 0.2885.79 ± 0.116<0.001
trans-Rose oxide11260.76 ± 0.1321.88 ± 0.038<0.001
Camphor1145nd0.25 ± 0.003-
Menthone11530.08 ± 0.0400.16 ± 0.0070.132
Isomenthone11645.78 ± 0.21810.61 ± 0.217<0.001
Citronellol122736.69 ± 1.57724.25 ± 0.495<0.01
Neral (β cis Citral)12400.27 ± 0.0280.00 ± 0.000<0.001
Carvone1244nd0.74 ± 0.015-
Geraniol125315.45 ± 1.69711.13 ± 0.142<0.05
Geranial12710.86 ± 0.0200.44 ± 0.009<0.001
Citronellyl formate127513.29 ± 0.21214.11 ± 0.2900.085
p-Menth-1-en-9-ol 12990.09 ± 0.0430.70 ± 0.015<0.001
Geranyl formate13024.21 ± 0.3534.75 ± 0.0990.215
Citronellyl acetate13520.21 ± 0.015nd-
Geranyl acetate13830.39 ± 0.048nd-
β-Bourbonene 13860.97 ± 0.160nd-
β-Caryophyllene14250.35 ± 0.009nd-
Citronellyl propanoate14500.35 ± 0.009nd-
Geranyl proponoate14870.86 ± 0.0990.49 ± 0.012<0.05
Germacrene D14971.35 ± 0.0600.34 ± 0.009<0.001
Viridiflorene15090.88 ± 0.020nd-
δ-Cadinene15340.45 ± 0.0310.42 ± 0.0090.354
Citronellyl butanoate15370.51 ± 0.018nd-
Geranyl butanoate15650.60 ± 0.0840.89 ± 0.020<0.05
Phenethyl tiglate15881.90 ± 0.1652.50 ± 0.049<0.05
γ-Eudesmol16216.92 ± 0.0795.92 ± 0.122<0.01
β-Εudesmol16510.39 ± 0.0210.64 ± 0.015<0.001
Citronellyl tiglate16650.44 ± 0.083nd-
Geranyl tiglate17002.43 ± 0.0521.58 ± 0.019<0.001
Farnesyl acetone2005nd1.22 ± 0.023-
Total Identified 99.65 ± 0.20899.75 ± 0.1280.713
Monoterpenes hydrocarbons 0.23 ± 0.0157.97 ± 0.364<0.001
Sesquiterpenes hydrocarbons 4.01 ± 0.1310.75 ± 0.014<0.001
Oxygenated monoterpenes 76.20 ±0.50873.04 ± 0.620<0.05
Oxygenated sesquiterpenes 10.19 ± 0.0328.13 ± 0.104<0.001
Others 9.03 ± 0.3899.85 ± 0.0950.111
nd: not detected.
Table 8. Chemical composition (%) of essential oils of rosemary plants grown at different altitudes (plain vs. mountain). Data are expressed as means ± SE (n = 3), and significant differences (p < 0.05, p < 0.01, p < 0.001) in bold expressed between treatments are indicated by Student’s t-test p-values.
Table 8. Chemical composition (%) of essential oils of rosemary plants grown at different altitudes (plain vs. mountain). Data are expressed as means ± SE (n = 3), and significant differences (p < 0.05, p < 0.01, p < 0.001) in bold expressed between treatments are indicated by Student’s t-test p-values.
ComponentsPlainMountainStudent’s t-Test
Tricyclene9220.22 ± 0.0030.20 ± 0.006<0.05
α-Thujene9260.01 ± 0.0030.04 ± 0.0340.341
α-Pinene93312.01 ± 0.25513.05 ± 0.6250.196
Camphene9488.29 ± 0.1358.12 ± 0.2730.614
β-Pinene9771.67 ± 0.021.71 ± 0.5130.942
n-Octanone9840.01 ± 0.0070.01 ± 0.0070.519
β-Myrcene9891.07 ± 0.0071.18 ± 0.1340.460
3-Octanol10030.03 ± 0.0000.03 ± 0.0301.000
α-Phellandrene10040.09 ± 0.0000.14 ± 0.0300.146
α-Terpinene10170.32 ± 0.0000.53 ± 0.1500.234
p-Cymene10243.03 ± 0.0613.03 ± 0.2480.990
Limonene10283.84 ± 0.0503.94 ± 0.1450.537
1,8-Cineole103132.94 ± 0.70332.94 ± 0.9270.996
γ-Terpinene10580.23 ± 0.0180.63 ± 0.3250.286
Terpinolene10890.22 ± 0.0070.38 ± 0.0840.118
Linalool11000.70 ± 0.0430.63 ± 0.1870.734
β-Thujone11160.02 ± 0.0090.09 ± 0.0350.100
Camphor114520.86 ± 0.40119.21 ± 0.445<0.05
Borneol11668.94 ± 0.1437.88 ± 1.0710.385
Terpinen-4-ol11780.92 ± 0.0350.97 ± 0.0410.465
p-Cymen-8-ol11850.06 ± 0.0060.03 ± 0.0180.224
α-Terpineol11913.05 ± 0.1072.58 ± 0.141<0.05
Bornyl acetate12850.56 ± 0.1030.85 ± 0.2540.355
Methyl eugenol14040.19 ± 0.0350.19 ± 0.0780.941
β-Caryophyllene14250.46 ± 0.0591.14 ± 0.212<0.05
α-Caryophyllene 14620.03 ± 0.0130.12 ± 0.012<0.01
δ-Cadinene 15340.03 ± 0.0090.10 ± 0.017<0.05
Caryophyllene oxide15870.06 ± 0.0150.04 ± 0.0260.678
α-Bisabolol16850.05 ± 0.0250.09 ± 0.0190.238
Total Identified 99.87 ± 0.05099.87 ± 0.0300.915
Monoterpenes hydrocarbons 30.97 ± 0.40932.96 ± 1.7740.335
Sesquiterpenes hydrocarbons 0.51 ± 0.0781.36 ± 0.239<0.05
Oxygenated monoterpenes 67.48 ±0.23364.33 ± 2.1510.219
Oxygenated sesquiterpenes 0.11 ± 0.0390.13 ± 0.0440.638
Others 0.79 ± 0.1331.07 ± 0.3680.519
Table 9. Chemical composition (%) of essential oils of sage plants grown at different altitudes (plain vs. mountain). Data are expressed as means ± SE (n = 3), and significant differences (p < 0.05, p < 0.01, p < 0.001) in bold between treatments are indicated by Student’s t-test p-values.
Table 9. Chemical composition (%) of essential oils of sage plants grown at different altitudes (plain vs. mountain). Data are expressed as means ± SE (n = 3), and significant differences (p < 0.05, p < 0.01, p < 0.001) in bold between treatments are indicated by Student’s t-test p-values.
ComponentsRIPlainMountainStudent’s t-Test
Salvene8650.11 ± 0.015nd-
Tricyclene9220.23 ± 0.0120.23 ± 0.0320.926
α-Thujene9260.18 ± 0.0060.13 ± 0.0670.525
α-Pinene9334.33 ± 0.1335.62 ± 0.479<0.05
Camphene9487.10 ± 0.2588.13 ± 0.4930.138
β-Pinene9772.90 ± 0.1013.31 ± 0.2270.179
β-Myrcene9891.69 ± 0.1101.47 ± 0.0430.136
α-Terpinene10170.12 ± 0.0090.05 ± 0.0500.222
o-Cymene10241.42 ± 0.1150.71 ± 0.052<0.01
Limonene10282.15 ± 0.0592.46 ± 0.055<0.05
1,8-Cineole103116.69 ± 0.52415.28 ± 0.3590.091
γ-Terpinene10580.20 ± 0.0210.29 ± 0.010<0.05
Terpinolene10890.03 ± 0.0300.18 ± 0.021<0.05
Linalool11000.14 ± 0.0030.06 ± 0.0570.202
α-Thujone110623.83 ± 0.0715.34 ± 0.426<0.001
β-Thujone11165.23 ± 0.17613.32 ± 0.665<0.001
iso-3-Thujanol1133nd0.33 ± 0.031-
trans-Sabinol11400.30 ± 0.0180.47 ± 0.006<0.001
Camphor114522.26 ± 1.02516.98 ± 0.474<0.01
Borneol11663.54 ± 0.12513.01 ± 0.656<0.001
Terpinen-4-ol11780.81 ± 0.0490.41 ± 0.023<0.001
α-Terpineol11910.23 ± 0.0240.15 ± 0.0780.417
Estragol11970.16 ± 0.009nd-
Bornyl acetate12850.59 ± 0.0492.21 ± 0.421<0.05
trans-sabinyl acetate12920.11 ± 0.009nd-
Copaene13490.02 ± 0.0230.41 ± 0.067<0.01
β-Caryophyllene14250.48 ± 0.1413.49 ± 0.360<0.001
α-Caryophyllene14620.86 ± 0.1790.48 ± 0.0520.111
γ-Cadinene1525nd0.29 ± 0.047-
δ-Cadinene1534nd0.71 ± 0.115-
Caryophyllene oxide15870.15 ± 0.0330.27 ± 0.0350.062
Viridiflorol15942.77 ± 0.2970.88 ± 0.032<0.01
Humulene epoxide II16080.64 ± 0.0650.00 ± 0.000<0.001
Cubenol1643nd0.42 ± 0.069-
neo-5-Cedranol 1699nd0.66 ± 0.174-
Manool20550.72 ± 0.1842.25 ± 0.514<0.05
Total Identified 99.97 ± 0.030100.00 ± 0.0000.374
Monoterpenes hydrocarbons 20.35 ± 0.42622.58 ± 1.2840.174
Sesquiterpenes hydrocarbons 1.36 ± 0.3045.38 ± 0.572<0.01
Oxygenated monoterpenes 73.20 ± 0.36865.53 ± 0.228<0.001
Oxygenated sesquiterpenes 3.55 ± 0.3242.23 ± 0.270<0.05
Others 1.53 ± 0.1784.45 ± 0.840<0.05
nd: not detected.
Table 10. Chemical composition (%) of essential oils of spearmint plants grown at different altitudes (plain vs. mountain). Data are expressed as means ± SE (n = 3), and significant differences (p < 0.05, p < 0.01, p < 0.001) in bold between treatments are indicated by Student’s t-test p-values.
Table 10. Chemical composition (%) of essential oils of spearmint plants grown at different altitudes (plain vs. mountain). Data are expressed as means ± SE (n = 3), and significant differences (p < 0.05, p < 0.01, p < 0.001) in bold between treatments are indicated by Student’s t-test p-values.
ComponentsRIPlainMountainStudent’s t-Test
α-Pinene9331.01 ± 0.0520.81 ± 0.040<0.05
Camphene9480.08 ± 0.0060.09 ± 0.0060.288
Sabinene9730.67 ± 0.0321.26 ± 0.061<0.001
β-Pinene9771.32 ± 0.0460.88 ± 0.3800.311
β-Myrcene9890.63 ± 0.0320.55 ± 0.0260.123
3-octanol9950.15 ± 0.0090.30 ± 0.024<0.01
Limonene102812.07 ± 0.3026.65 ± 0.055<0.001
1,8-Cineole10314.98 ± 0.2345.68 ± 0.150<0.05
cis-β-Ocimene 10360.13 ± 0.0060.18 ± 0.009<0.05
trans-β-Ocimene 1046nd0.06 ± 0.003-
γ-Τerpinene10580.04 ± 0.0030.09 ± 0.003<0.001
cis-Sabinene hydrate10670.19 ± 0.0120.30 ± 0.017<0.01
3-Octanol acetate1121nd0.10 ± 0.003-
iso-Μenthone11640.08 ± 0.003nd-
Borneol11660.26 ± 0.0120.25 ± 0.0090.670
Terpinen-4-ol11780.12 ± 0.0090.23 ± 0.012<0.001
α-Τerpineol11910.19 ± 0.0090.07 ± 0.006<0.001
cis-Dihydro carvone11980.65 ± 0.03312.97 ± 0.613<0.001
neo-Dihydro carveol11940.38 ± 0.0171.34 ± 0.042<0.001
trans-Carveol1219nd0.29 ± 0.015-
cis-Carveol12310.13 ± 0.0093.60 ± 0.159<0.001
Pulegone12400.70 ± 0.0260.44 ± 0.020<0.001
Carvone124472.12 ± 0.91150.12 ± 1.203<0.001
cis-Carvone oxide12620.07 ± 0.006nd-
trans-Carvone oxide12760.13 ± 0.006nd-
iso-Bornyl acetate1285nd0.08 ± 0.003-
Dihydrocarveol acetate13250.43 ± 0.0205.67 ± 0.278<0.001
trans-Carvyl acetate1335nd0.24 ± 0.012-
cis-Carvyl acetate 13600.27 ± 0.0154.92 ± 0.187<0.001
β-Bourbonene13860.61 ± 0.0190.61 ± 0.0191.000
β-Elemene b13930.21 ± 0.0090.19 ± 0.0280.539
β-Caryophyllene14250.92 ± 0.0680.91 ± 0.0310.900
cis-Muurola-3,5-diene 14560.42 ± 0.0200.42 ± 0.0181.000
Germacrene D14970.38 ± 0.0230.31 ± 0.0240.116
Bicyclogermacrene15120.16 ± 0.0060.10 ± 0.0380.179
Germacrene A15190.07 ± 0.0030.07 ± 0.0120.621
trans-Calamene 15310.24 ± 0.0150.20 ± 0.0070.105
1,10-di-epi Cubenol 16420.10 ± 0.000nd-
Total Identified 99.91 ± 0.003100.00 ± 0.000<0.001
Monoterpenes hydrocarbons 15.95 ± 0.42910.58 ± 0.463<0.001
Sesquiterpenes hydrocarbons 3.01 ± 0.1632.81 ± 0.0870.034
Oxygenated monoterpenes 79.59 ± 0.64074.03 ± 0.360<0.01
Oxygenated sesquiterpenes 0.10 ± 0.000nd-
Others 1.23 ± 0.05712.56 ± 0.076<0.001
nd: not detected.
Table 11. Correlations between antioxidant activity, phenolic compounds, and essential oil components.
Table 11. Correlations between antioxidant activity, phenolic compounds, and essential oil components.
Activity Assay
PhenolsFlavonoidsFlavanolsEssential Oils (ΕO)1,8 CineoleCamphorBorneolcis-Dihydro AgarofuranSilphiperfol-5-En-3-One A
DPPH + +
PhenolsFlavonoidsFlavanolsEOIsomenthoneCitronellolGeraniolCitronellyl formateγ-Eudesmol
PelargoniumPFRAP ++
PhenolsFlavonoidsFlavanolsEOα-PineneSabineneβ-Pinene1,8-CineoleTerpinyl acetate a
LaurelPFRAP+++ +-
DPPH+++ -
PhenolsFlavonoidsFlavanolsEO1,8 CineoleLinaloolCamphorBorneolCarvone
PhenolsFlavonoidsFlavanolsEOD-Limonene1,8-CineoleNeralGeranialCaryophyllene oxide
Lemon verbenaPFRAP +
MFRAP++ -+---
PhenolsFlavonoidsFlavanolsEOD-limonene1,8-Cineolecis-dihydro carvoneCarvoneDihydrocarveol acetate
The plus (+) and minus (−) symbols indicate positive and negative correlations, respectively. The P and M indicate the plain and mountain areas, respectively.

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Chrysargyris, A.; Mikallou, M.; Petropoulos, S.; Tzortzakis, N. Profiling of Essential Oils Components and Polyphenols for Their Antioxidant Activity of Medicinal and Aromatic Plants Grown in Different Environmental Conditions. Agronomy 2020, 10, 727.

AMA Style

Chrysargyris A, Mikallou M, Petropoulos S, Tzortzakis N. Profiling of Essential Oils Components and Polyphenols for Their Antioxidant Activity of Medicinal and Aromatic Plants Grown in Different Environmental Conditions. Agronomy. 2020; 10(5):727.

Chicago/Turabian Style

Chrysargyris, Antonios, Maria Mikallou, Spyridon Petropoulos, and Nikolaos Tzortzakis. 2020. "Profiling of Essential Oils Components and Polyphenols for Their Antioxidant Activity of Medicinal and Aromatic Plants Grown in Different Environmental Conditions" Agronomy 10, no. 5: 727.

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