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Article

Arbuscular Mycorrhizal Fungi (AMF) Influence Yield and Essential Oil Content and Composition of Sage (Salvia officinalis L.) Under Different Water Regimes

by
Andrzej Sałata
1,
Renata Nurzyńska-Wierdak
1,*,
Andrzej Kalisz
2,
Sara Ibánez-Asensio
3,
Héctor Moreno-Ramón
3,
Joanna Majkowska-Gadomska
4 and
Anna Francke
4
1
Department of Vegetable and Medicinal Plants, Faculty of Horticulture and Landscape Architecture, University of Life Sciences in Lublin, 20-950 Lublin, Poland
2
Department of Horticulture, Faculty of Biotechnology and Horticulture, University of Agriculture in Krakow, 31-425 Krakow, Poland
3
School of Agricultural Engineering and Environment, Universitat Politècnica de València, 46022 València, Spain
4
Department of Agroecosystems and Horticulture, Faculty of Agriculture and Forestry, University of Warmia and Mazury in Olsztyn, Plac Łódzki 3, 10-718 Olsztyn, Poland
*
Author to whom correspondence should be addressed.
Agronomy 2025, 15(12), 2753; https://doi.org/10.3390/agronomy15122753
Submission received: 4 November 2025 / Revised: 24 November 2025 / Accepted: 27 November 2025 / Published: 28 November 2025
(This article belongs to the Special Issue Cultivation and Utilization of Herbal and Aromatic Plants)
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Abstract

Essential oil-bearing plants are valued for their aromatic qualities and medicinal value. An example of such a plant is sage (Salvia officinalis L.), one of the most important aromatic herbal plants. Rich in essential oil (EO), sage herb is used in pharmaceutical and cosmetic production and as a spice. This study was conducted to determine the effect of arbuscular mycorrhizal fungi (AMF) on the morphological characteristics, yield, and EO production of sage under different irrigated conditions: 95 ± 5, 75 ± 5, 50 ± 5, and 25 ± 5% field capacity (FC). Maximum herb dry biomass yield and leaf dry biomass yield were obtained at the 95, 75% FC irrigation level, and the highest leaf EO (1.361%) was at 75% FC. The high yield of herb and leaf dry biomass, as well as the highest EO in herb and leaves, was observed with AMF. A group of monoterpenes and sesquiterpenes dominated the EO. The main compounds were 1,8-cineole, α-thujone, β-thujone, camphor, E-caryophyllene, and viridiflorol. The chemical composition of EO has changed under irrigation. Under severe stress (25% FC) and moderate soil moisture (50% FC), the proportion of monoterpene hydrocarbons and oxidized sesquiterpenes was higher than at 75% and 95% FC. The proportion of camphor, α-thujone, and β-thujone was higher with AMF only under severe water stress. With increasing soil moisture, the proportion of α-thujone and β-thujone in EO decreased. A positive correlation was found between EO content and the number of leaves per plant. Our results indicate the prospects for practical application of AMF in combination with the irrigation of sage plants.

1. Introduction

A modern plant drug is a product defined in terms of the chemical compounds biosynthesized by plants. The pharmaceutical properties of sage (Salvia officinalis L.) are determined by secondary metabolites, such as essential oil (EO) constituents, including terpenoids like monoterpenes, sesquiterpenes, and their oxygen derivatives, phenylpropanoids, also phenolic acids, and flavonoids [1,2]. The main components of sage EO are α- and β-thujone (usually accounting for 50%), camphor, cineole, and borneol [3,4]. EO is secreted and accumulated in specialized glandular trichomes of the plant [5]. Sage EO exhibits high antioxidant activity and antimicrobial activity, especially against Bacillus subtilis [6,7]. EO is used in oncology and for the treatment of neurodegenerative diseases [8,9]. Several studies have demonstrated the anti-diabetic, anti-inflammatory, and memory-enhancing effects of sage and its constituents [10]. Biomass harvested before sage blooms is widely used in the food industry as an additive to improve flavor [2]. Drought and soil salinity are major stress factors that reduce crop yields and can lead to permanent damage and even plant death [11]. For many years, scientists have paid much attention to droughts, especially those caused by rainfall deficiency. Amani Machiani et al. [12] found a reduction in the content and deterioration of the EO of Thymus vulgaris, and Li et al. [13] found a reduction in the EO of Salvia miltiorrhiza under water stress conditions.
From the point of view of yield biology, the introduction of mycorrhizal preparations into the substrate has a beneficial effect on plants: it increases the assimilation of hard-to-access components, improves humus-forming capacity, eliminates pathogens, and improves growth and yield quality [6]. Numerous studies report positive effects of AMF on increasing terpenoid levels in such medicinal plants like Salvia officinalis, T. daenensis, Ocimum tenuiflorum, and T. vulgaris [12,14,15]. AMF can be a factor mitigating abiotic stress and improving the growth and quality of plant crops under unfavorable conditions. Zhang et al. [16] and Sun and Shahrajabian [17] observed that, under drought conditions, mycorrhiza with plant roots enables the maintenance of a high leaf water potential. It is most often associated with a better-developed root system and more efficient use of soil water. Ectomycorrhizal fungi can support plant growth by reducing stress from water or phosphorus deficiency [18]. AM fungi are involved in nitrogen mobilization from organic compounds and increase nitrogen uptake from the soil by plants [19]. Zhang et al. [20] showed that through mycorrhiza, plant water management can adapt to increasing water scarcity by reducing transpiration, efficient water uptake by a well-developed root system, efficient conduction, or the ability to store water [21]. During stress, AMFs optimize plant growth and development while reducing energy expenditure [22]. One of the factors determining the biological activity of AM mycelium is the water level in the soil [23]. Peñuelas and Staudt [24] and also Konvalinková and Jansa [25] pointed out that soil water availability significantly affects mycorrhiza formation, mycelial activity, and diversity, ultimately affecting the synthesis of secondary metabolites in the plant.
The effect of irrigation on the content and composition of sage EOs has been studied under warm climate conditions [26]. In aromatic plants, the growth and production of EOs are often influenced by various environmental factors, including climate, soil, and sunshine [27], with soil water content being one of the key factors [28]. According to Karousou et al. [29], under very adverse conditions, the plant’s survival strategy is to drastically inhibit the very energy-intensive growth process. Also, the elimination of damage to cellular structures and organs, and the regeneration of the plant already after the stress factor has subsided, requires large amounts of energy [30]. It usually comes at the expense of lowering yield and also lowering EO content [31]. The benefits of irrigating herbaceous plants have been documented in the literature for a long time [32,33]. According to Blum [34], under water deficit conditions, there is a decrease in turgor, resulting in a decrease in both growth and cell development in the aboveground parts of sage, primarily stems and leaves. As demonstrated by Soltanbeigi et al. [35], the magnitude of the decrease in biomass and EO content under water deficit conditions depends on the severity and duration of water stress. Despite its beneficial effects on increasing biomass and yield EO, irrigation generally does not cause significant changes main constituents of EO [36,37]. According to Mossi et al. [38] and Grisafi et al. [39], sage has broad adaptability, exhibits drought and low temperature tolerance, and has a high capacity for biomass production and regeneration. According to Somnez and Bayram [28], in agricultural practice, irrigation combined with nitrogen fertilization significantly increases the yield of sage but causes a reduction in the quantity and quality of EO in the herb, such as by reducing the antioxidant value. It has been noted [40,41] that with irrigation, the rate of mineral release from the sorption complex coincides optimally with the nutritional requirements of sage.
Most studies have focused exclusively on the impact of a single factor of drought stress [27,28]. Only two experiments of combined AMF inoculation and irrigation have been conducted. A study by Hamedani et al. [42] dealt with colonization of Sesamum indicum L. roots by two AMF species (i.e., Rhizophagus intraradices and Funneliformis mosseae), and a study by Khajeeyan et al. [43] concerned the inoculation of Aloe vera with Glomus mosae fungi. The functioning of symbiosis requires appropriate environmental conditions. The effectiveness of establishing mycorrhiza with plant roots may also change as the plants grow and go through developmental stages. The literature on the integrative application of AMF in combination with irrigation on the quantity and quality of EOs of aromatic plants is limited. Considering the results of the cited studies, the present study attempted to verify the research hypothesis: the productivity, morphological characteristics, and quantity and composition of sage EOs depend on the mycorrhizal activity of AMF at different irrigation levels. The purpose of the research undertaken was (i) to identify the sensitivity of sage to soil water deficit stress and its alteration by AMF treatment, (ii) to evaluate the effect of varying soil moisture and AMF on yield and selected morphological characteristics, and (iii) to determine the effect of different levels of water in the soil and fungi inoculation on the content and composition of sage EO.

2. Materials and Methods

2.1. The Study Site and Soil Parameters

Agrotechnical experiment was conducted at the research station of the University of Life Sciences in Lublin (UP), located in southeastern Poland (51.23° N, 22.56° E). Chemical composition analyses were made at the Central Laboratory of UP in Lublin. The experiment was conducted from May to September 2021 in an unheated high tunnel covered by 200 µm (200 g m−2) thick, single-layer, and clear polyethylene film. The soil in the tunnel was clay-sandy [44], characterized by the following composition (0–20 cm): clay (<0.002 mm) 9%, sand (2–0.05 mm) 25%, dust (0.05–0.002 mm) 66%, and pHKCl-6.5 (potentiometrically). Nutrient element contents were as follows (g kg−1): N 1.4; C 19.1; and (in mg kg−1): phosphorus 16.5, potassium 220, and magnesium 9. The average external daily temperature during the growing season (May–September) was 17.4 °C; maximum temperature 22.0 °C; minimum temperature 17.1 °C; relative humidity: 55%; total sunshine hours were 968.

2.2. Plant Material and Field Management

The experimental material consisted of sage (Salvia officinalis L.) plants, whose seeds were obtained from the Polish seed company PNOS Ożarów Mazowiecki. Sage seedlings were produced in the greenhouse. Seeds were sown in the first decade of April 2021 into multi-pots filled with peat substrate, Hartmann Poland (the capacity of a single pot was 90 cm3). The plants were fertilized twice with a 0.1% solution of Florovit multicomponent fertilizer (Inco S.A., Warsaw, Poland), with a content of 3.0% nitrogen (2.3% amide nitrogen and 0.7% nitrate nitrogen), 2.4% K2O, 0.006% copper, 0.03% iron, 0.015% manganese, 0.0017% molybdenum, and 0.013% zinc, at pH 3.3–4.3. Well-developed (10 cm high and four pairs of leaves) and hardened-off sage plants were planted into the ground in the tunnel on May 5.

2.3. Experimental Design

2.3.1. Setup of the Experiment

The experiment was conducted in a dependent sub-block arrangement (split-plot) with three replications. Sage plants were grown at a spacing of 0.4 × 0.4 m (40 plants in each replication on a plot of 6.4 m2). The following experimental factors were applied in the experiment: Factor I (irrigation): at 95 ± 5, 75 ± 5, 50 ± 5, and 25 ± 5% of field capacity (FC); factor II (mycorrhiza): arbuscular mycorrhizal fungi (AMF); control object without mycorrhiza (non-AMF). There were a total of 24 experimental plots in the experiment.

2.3.2. Preparation and Inoculation of Arbuscular Mycorrhiza

Mycorrhization was performed with ectomycorrhizal symbiotic mycelium. Their producer and distributor was the Mycorrhizal Fungi Laboratory Mykoflor® (Końskowola, Poland). The isolate contained spores and mycelia of mycorrhizal fungi (Rhizophagus aggregatus, R. intraradices, Claroideoglomus etunicatum, Endogone mosseae, Funneliformis caledonium, and Gigaspora margarita). The manufacturer recommends mycelium for all vegetable and herbaceous plants except those in the Brassicaceae family. Inoculation was carried out according to the manufacturer’s recommendations on a one-time basis, with a dose of 3 mL, or 100 propagation units. Each propagation unit can colonize one portion into the root zone. The AMF solution was prepared by dissolving 1 g of dry inoculate in 300 mL of distilled water. According to the certificate presented by the manufacturer (Mycoflor®), 1 g of dry inoculate contains approximately 10,000 colony-forming units (CFUs) of live propagules. The mycelium contained an added hydrogel, a dose of moisture necessary for the mycelium during the first period of development.

2.3.3. Irrigation

Irrigation treatments started two weeks after the seedlings were entirely rooted in the soil. Soil moisture content was determined using the weight (dryer-weight) method at each irrigation treatment date by measuring the difference between the weight of a wet soil sample and the weight of the same sample dried at 105 °C for 24 h. The relationship between water potential and soil moisture at specific points was determined using a soil moisture characteristic curve. To control soil water potential, continuous measurement was carried out with a tensiometer (Irrometer Company Inc., Riverside, CA, USA). Water doses and irrigation frequency were adjusted to the optimum soil moisture level and optimum irrigation time. Plants were irrigated using an in-ground system with T-Tape Rivulis 508-20-400 drip lines (Milex, Dobrzyków, Poland), set between rows every 40 cm, at a distance of 3–4 cm from the plants. Foils were placed between the sites with different irrigation levels (95, 75, 50, and 25% FC) to a depth of 1 m to eliminate lateral water sucking. During periods of water scarcity, a drip line with a capacity of 4.0 L m h−1, with an operating pressure of 1.5 bar, was used. In all irrigation treatments, nine irrigations were carried out. A total of 3940, 3400, 1500, and 1380 m3 ha−1 of water were used for irrigation treatments of 95, 75, 50, and 25% FC, respectively.

2.3.4. Other Agronomic Practices

Two weeks before planting the seedlings, a multi-nutrient (NPK 4-3-3 + 1 MgO BIO) organic fertilizer, Fertikal (Beveren, Belgium), at a rate of 0.1 kg per 1 m2, was applied. The fertilizers were mechanically mixed into the soil to a depth of about 5 cm. During the growing period, the plants were fertilized twice with nitrogen in the form of ammonium nitrate with a concentration of 34% N (the single dose was about 5 kg N ha−1), and the necessary cultivation treatments were carried out (hand weeding several times, loosening the soil in the inter-rows). No chemical pesticides were used during the cultivation period, and no diseases or pests were observed in the crop.

2.3.5. Harvest and Plant Measurements

Sage herb was harvested from annual plants (irrigated, with AMF and non-AMF) once on September 10. Sage herb (raw leafy stems with inflorescences) was harvested at the beginning of plant flowering by hand using a knife, cutting the herb 5 cm above the soil surface above the first branching. The yield of fresh herb was calculated based on the weight of stems with leaves and inflorescences (kg m−2) and converted to an area of 1 ha. Seedling height was measured from the base of the stem of the main shoot to the tip of the longest leaf, and the number of branches on the plant was counted. All leaves longer than 3 cm were counted from individual plants. Measurements were made on 10 plants in each combination. Immediately after harvesting, samples were prepared for chemical analysis and for drying separately from irrigated, AMF, and non-AMF plants. The convective drying process was carried out in a drying room with a 35 °C air stream flowing parallel to the layer at a speed of 0.5 m s−1, in total darkness. The weight of the herb during drying was 2.0–2.5 kg m−2 area. Drying of the raw material consisted of gradually increasing the temperature by 5 °C to a final temperature of 35 °C with fans open. Conditioning took 24 h to remove residual water, with the fans closed. After drying, the raw material contained 12–14% water in five consecutive measurements. The yield of air-dried herb was calculated (g m−2), and the herb was then grated on sieves to separate the leaves from the stems to determine the yield of grated leaves (g m−2). After conversion, the yield of fresh herb, the yield of air-dried herb, and the yield of air-dried leaves were expressed in units of t ha−1. Samples of 0.25 kg were taken from air-dried plant material and stored in airtight containers until laboratory testing. The content of EO (%) was determined in fresh biomass, separately for the herb and leaves.

2.4. Chemical Analyses

2.4.1. Distillation of EO

The air-dried leaves, after weighing the samples (20 g each), were placed in 1 L glass flasks, flooded with 400 mL of water, and destined for distillation conducted in a Clevenger-type apparatus for 3 h, starting from the moment the flask contents boiled and the first drop was collected. The intensity of heating was adjusted so that 3–4 mL of liquid per minute flowed into the receiver. After the distillation was completed, cooling was turned on, the oil was brought to the microscale, and the result was read after 30 min.

2.4.2. EO Composition

The quantitative and qualitative composition of EO obtained from air-dried sage leaves was determined by gas chromatography-mass spectrometry (GC-MS) on a Varian 4000 MS/MS apparatus with a VF-5 m column (DB-5 equivalent). A measurement range of 40–1000 m/z and a scan rate of 0.8 s/scan were used. The carrier gas was helium, and the constant flow rate was 0.5 mL min−1. The dispenser temperature was 250 °C, and a temperature gradient of 50 °C was applied for 1 min, then increased to 250 °C at a rate of 4 °C min−1 and maintained at 250 °C for 10 min. Split 1:1000 m/z, 1 μL of solution was dispensed (10 μL of sample in 1000 μL of hexane). Non-isothermal Kovacs retention indices were determined based on the range of C10-C40 alkanes. Qualitative analysis was based on the NIST/EPA/NIH Mass Spectral Library [45]. The identity of the compounds was confirmed based on their retention indices taken from the literature [46] and our data.

2.5. Statistical Elaboration

The normality of the distribution of the studied traits was checked using the Shapiro–Wilk W test (p < 0.05). A two-factor analysis of variance (ANOVA) was used to test significance levels at p ≤ 0.05, p ≤ 0.01, and p ≤ 0.001 for plant height, number of leaves and branches, yield of fresh and air-dry herb, leaf biomass, and EO total content and components that were most significant for EO quality (1,8-cineole, α-thujone, β-thujone, camphor, E-caryophyllene, viridiflorol). A post hoc least significant difference (LSD) test was used to separate means into homogenous groups. Tukey’s test confirmed significant differences between factors at above-mentioned significance levels. Relationships between plant morphological traits, yield of fresh and air-dry herb and leaf biomass, and EO content and composition were estimated using Pearson’s correlation coefficients [47]. Data analysis was performed using Statistica PL software, version 13.0 (StatSoft Inc., Tulsa, OK, USA).

3. Results

3.1. Morphological Traits of Plants

The irrigation and AMF levels used significantly affected plant height, number of leaves, and number of branches per plant (Table 1). At irrigation levels of 75% and 95% FC, plants were taller by an average of 5 cm, produced an average of 10 more leaves, and formed more branches than under soil water deficit conditions (25% and 50% FC). More than 12 leaves were produced by plants grown with AMF than without AMF. There was no significant effect of AMF on the height and number of branches per plant.
Under soil water deficit conditions (50% FC and 25% FC), the number of leaves per plant was significantly higher in the crop with AMF compared to the crop without AMF (Figure 1). There was no significant effect of AMF under conditions at 95% and 75% FC irrigation levels.

3.2. Yield of Fresh and Air-Dry Biomass

The total yield of fresh and air-dry biomass at irrigation levels of 75 and 95% FC was 18% higher, and the total biomass of fresh and air-dry leaves was 36–38% higher compared to soil water deficit conditions (25–50% FC) (Table 1 and Table 2). In the crop with AMF, dry biomass yield was 397 kg ha−1, and dry leaf yield was 250 kg ha−1, which were 98 and 73 kg ha−1 higher, respectively, than without AMF. AM fungi did not affect fresh biomass. A significant interaction of irrigation and AMF treatment was confirmed for dry herbage yield and air-dry leaf yield (Figure 2). At each irrigation level (25%, 50%, 75%, and 95% FC), dry biomass yield and dry leaf yield were higher in crops with AMF than without AMF.

3.3. EO Content

In the herb at irrigation levels of 50%, 75%, and 95% FC, the average EO content was 0.771–0.862%, which was 24–32% higher than under strong soil water deficit conditions (25% FC) (Table 2). The EO content of sage leaves was higher at the irrigation level of 75% FC, and it was 1.361%. This amount was higher by 16% (50% FC), by 27% (25% FC), and by 34% (95% FC) than at a low soil water level (25% FC). The high calculated EO yield was 28.55 kg ha-1 (75% FC), which was 10–15 kg ha−1 higher than at the 25%, 50%, 95 percent FC levels. In the crop with AMF, the EO content in the herb was 0.950%, and in the leaves, it was 1.306%, which was 42% and 32% higher, respectively, compared to the crop without AMF. The yield of EO in the crop with AMF was 9 kg ha−1 higher than that without AMF. By far the most significant amount of EO was accumulated in the herb and leaves in the crop with AMF with irrigation at 50% FC (1.192% and 1.704%, respectively) (Figure 3). Compared to the crop without AMF, EO content in the herb in the crop with AMF was higher at 50% and 25% FC, and in the leaves at 50% and 75% FC. EO yield was highest in the crop with AMF and irrigation levels of 75 and 50% FC (29.63 and 26.35 kg ha−1), respectively, and lower without AMF at 50% FC (10.43 kg ha−1) (Figure 4).

3.4. EO Composition

The sage EO studied was dominated by a group of monoterpenes and sesquiterpenes (Table 3). The dominant compounds were α-thujone and camphor (Table 4). The chemical composition of EO changed under irrigation. Under water-deficient conditions (25% FC), the proportion of monoterpene hydrocarbons (MH) from both AMF and non-AMF was higher than at 50–75% FC. The proportion of oxidized sesquiterpenes (OS), but only for non-AMF, was higher at soil moisture content of 75% FC and under water-deficient and medium water content conditions (25 and 50% FC). On the contrary, at soil moisture of 75 and 95% FC, the proportion of oxidized monoterpenes (OM) from non-AMF only and hydrogenated sesquiterpenes (SH) from AMF was higher.
Under severely stressed conditions (25% FC), there were more α-thujone and β-thujone (21.229% and 8.077%, respectively). The share of E-caryophyllene was higher at 50 and 25% FC, and viridiflorol at 50% FC (Table 4). Under conditions of optimal soil moisture (75% FC), the share of 1,8-cineole and camphor was by far the highest, while at the same time, the lowest levels of α-thujone, β-thujone, E-caryophyllene, and viridiflorol were recorded at 75% FC. There was no significant effect of mycorrhiza on α-thujone, β-thujone, camphor, and viridiflorol. Cultivation with AMF caused higher accumulation of 1,8-cineole and E-caryophyllene (by 10% and 26%, respectively) than without AMF. A significant interaction ‘irrigation AMF’ was confirmed for 1,8-cineole, α-thujone, β-thujone, camphor, and E-caryophyllene (Figure 4, Figure 5 and Figure 6). The proportion of 1,8-cineole and E-caryophyllene in EO with AMF was higher at all water content levels than without AMF. The content of α-thujone and β-thujone was higher for AMF treatment, mainly under 75% FC. In contrast, the proportion of camphor was higher in the crop with AMF under water stress conditions (25% FC).

3.5. Correlation Analysis

Plant height, the number of leaves per plant, and the degree of branching of the plant were correlated with yield of herb biomass, leaf biomass, air-dry herb biomass, and air-dry leaf biomass (Figure 7). There was a correlation (0.59) between the number of leaves and EO content and EO yield. There was no significant correlation between 1.8-cineole, as the most characteristic compound of sage EOs, and morphological characteristics and EO content of the herb or leaves. However, a positive correlation was observed between the proportion of α-thujone and its isomer β-thujone (<5%) in sage EOs.

4. Discussion

4.1. Morphological Characteristics and Biomass Yield as Affected by Irrigation

Irrigation largely determined plant morphological traits and yield. At water deficit (50% and 25% FC), plant height, number of leaves, and branches were lower. This result was consistent with an earlier study by Sezen et al. [48], who found that, under water stress conditions, Salvia splendens plants produced fewer branches and lower biomass. Water deficiency reduces solar energy absorption and nutrient uptake by reducing the number and area of leaves [49]. The measured biological parameters that determine plant productivity are total fresh weight and dry weight. Dry mass is a better indicator of changes in plant biomass than fresh mass because it does not take into account water content, which can vary with growing conditions. In this work, the dry weight of the aboveground parts of sage and leaves showed similar trends for different soil water levels. At irrigation levels of 95 and 75% FC, the yield of air-dry biomass was 45% higher, and that of sage leaves was 60% higher compared to 50 and 25% FC. The positive effect of water at 50–75% FC on biomass growth of S. officinalis was found by other research teams, Rioba et al. [41] and Soltanbeigi et al. [35].

4.2. Effect of Irrigation on Oil Quantity and Quality

EO content in sage herb ranged from 0.585% to 0.862%, in leaves from 0.894% to 1.361%, depending on irrigation level. The amount of EO in sage herb was similar to the results of Zawiślak [50], but lower than the amount of 0.4–2.5% (% g/g) reported by Maksimovic et al. [51], which is most likely due to ontogenetic and environmental diversity. More sage EO was found at irrigation levels of 75 and 50% FC, indicating that moderate soil water content increases its amount. However, high water deficiency (25% FC) versus 75% FC reduced EO levels in leaves by 38%, in the herb by 47%, and EO yield by 68%. Additionally, a high water level in the soil (95% FC) was responsible for a significant reduction in EO levels in the herb by 52%, resulting in a 47% decrease in EO yield. Under water-deficient conditions (25% FC), 1,8-cineole and camphor were lower, and α-thujone, β-thujone, and E-caryophyllene were higher compared to 75% FC. Mohammadi et al. [52] explained that the content of EOs decreases under water stress due to a decrease in the activity of enzymes and biochemical pathways involved in their synthesis. According to Cal et al. [53] and Asghari et al. [54], under unfavorable environmental conditions, plants reduce the growth and synthesis of EOs, as this is an evolutionarily evolved and established mechanism by plants to avoid unnecessary energy inputs. This is confirmed by our research results. In this study, moderate stress led to an increase in the percentage of EO in plants due to the synthesis of terpenes and phenols. At 50% FC, the oil content in the herb was high and did not differ significantly from that at 75–95% FC, but oil yield was low.
There is no clear information in the scientific literature on the effect of irrigation or drought stress on EO content. For example, under controlled conditions, the highest amount of EO was observed after moderate drought stress (105 ± 5 mm evaporation) than regular (70 ± 5) mm evaporation [35]. In addition, moderate deficit increased the essential oil yield and the main constituents, camphor, α-thujone, and 1.8-cineole [26]. In contrast, García-Caparrós et al. [31] did not confirm that drought stress increased the accumulation or altered the composition of EOs in Mentha piperita, Salvia lavandulifolia, Thymus capitatus, and T. mastichina. The EO content only decreased in Lavandula latifolia and Salvia sclarea plants under water deficit conditions. These conflicting results may indicate interactions between plant response to soil moisture and other environmental/agronomic conditions. The quantity and quality of these secondary metabolites are affected by abiotic stress factors [55]. It can be posited that stresses typically act as a catalyst for the secondary metabolism of sage plants, resulting in increased production of EO compounds (e.g., 1,8-cineole, linalool, camphor, and borneol) and subsequent enhancement of their biological activities [56].

4.3. Effect of AMF on Plant Morphology and Biomass

AMF are sensitive to both low and excessive soil moisture. Excess water can adversely affect mycorrhizal development, as AMF are microorganisms sensitive to reduced oxygen availability. The drought resistance of mycorrhizal plants depends on the AMF species colonizing plant roots. It is assumed that MF can modify the growth of above-ground plant parts under drought stress, influencing the complex mechanisms of plant response to water deficit, related to P and K uptake, root respiration, photosynthesis, transpiration, and leaf osmotic potential [25,54]. The indicator for assessing AMF-plant symbiosis is the quantification of the rate of plant root colonization. Our research focused on the chemical analysis of the plant material and essential oil and did not take into account quantitative determinations on root colonization by AMF. The results showed that inoculation with AM fungi clearly affected the number of leaves. The effect of mycorrhiza was under soil water deficit conditions (moderate 50% and strong 25% FC), as plants with AMF developed more leaves. So, it can be indicated that in sage with AMF, there is a possibility of producing more leaves under drought conditions. However, many more leaves were on plants with AMF at high hydration levels (75% and 95% FC). The data obtained on the increase in the number of leaves as a result of mycorrhizal symbiosis can be explained by the increase in surface area of the roots due to the proliferation of the fungal hyphae [57]. So far, no studies have been conducted on the effectiveness of mycorrhiza in sage in relation to the water level in the soil [58]. However, Tarraf et al. [18] obtained a similar effect, increasing the number of leaves, by inoculating S. officinalis with AM fungi in combination with phosphorus.
The total dry herb biomass and leaf biomass of sage in mycorrhizal colonization were 25–29% higher than that of non-inoculated plants. At the same time, there was no significant effect of AMF on fresh herb and leaf biomass. Similarly, in research conducted by Nell et al. [59], there was no effect of inoculation (Symbivit, Glomus messeae, Glomus intraradices) on increasing the biomass of S. officinalis. On the contrary, in other studies [18], the biomass of S. officinalis leaves inoculated (Symbivit, Septoglomus viscosum) was 1.4–2.4 times higher than that of non-inoculated leaves. Previous studies [60] indicate that different species show differential interactions with the same AM.
Our results indicate (Figure 2) that, at each irrigation level, the yield of air-dry biomass of sage was higher with AMF than without mycorrhiza. It confirms the reciprocal relationship resulting from the functional activity of symbiosis between plant and fungus at varying levels of soil water content [61]. Apparently, the application of AMF improved plant growth parameters by reducing soil acidity and providing adequate conditions for the absorption of essential nutrients, especially nitrogen and phosphorus [18,62]. Reduced yields of sage herb and leaves at lower irrigation levels (50 and 25% FC) can be explained by lower availability of sufficient moisture around the root zone, and thus lower growth of root biomass. It leads to lower uptake of nutrients and water, harms photosynthesis rate, cell differentiation, and division, and ultimately reduces plant growth parameters and yield. Similarly, Amani Machiani et al. [61] noted that fresh yield and dry yield of thyme decreased under moderate (60% FC-by 15.1% and 13%, respectively) and severe water stress (40% FC–29.1% and 40.3%, respectively).

4.4. Effect of AMF on EO

Several potential mechanisms associated with terpenoid accumulation by AMF have been described in the literature, such as altering the levels of the phytohormones gibberellic acid (GA3), jasmonic acid (JA), and cytokine [63]. In most of these studies, increases in terpenoid content can be linked to phosphorus (P) availability and the transcription of genes responsible for terpenoid biosynthetic pathways [64,65]. The results of the experiment showed that AMF increased the amount of essential oil in sage herb and leaves. Comparable results were reported in S. officinalis by inoculation with Glomus viscosum fungi [18]. Application of AMF favorably improves nutrient balance, resulting in increased activity of enzymes and biochemical pathways involved in EO synthesis [66]. Similar results have been obtained in other medicinal plants, such as cilantro and basil [67,68]. However, it is unclear how AMFs increase the content of EOs in sage plants. For example, Binet et al. [69] showed no effect of AMs on EO content in Artemisia umbelliformis. Similarly, Tarraf et al. [70] did not confirm the effect of AMF colonization in T. vulgaris on EO accumulation. In a study by Amani Machiani et al. [61], the combined application of AMF and chitosan increased the amount of EO of thyme under water stress conditions. The authors explained that the association of AMF with thyme roots increases nutrient solubility and water uptake, while the application of chitosan reduces transpiration under stress conditions. AMF are among the most ubiquitous plant mutualists that enhance the growth and yield of plants by facilitating the uptake of nutrients and water [71].
No report was found on the effect of AMF application on sage EO at different irrigation levels. In our study, the highest EO in herb and leaves was from AMF under moderate moisture conditions at 50% FC irrigation levels. In herb and leaves with AMF, most EOs were with 50% FC irrigation level, and without AMF at 75%. In the crop with AMF, there were fewer EOs under both water deficit (25% FC) and high moisture conditions (75% FC). This shows that under moderate stress, AMF helps maintain the appropriate level of physiological and metabolic processes in the plant. In a study by Abdel-Salam et al. [72], limited soil water availability harmed root colonization of T. vulgaris by AMF. Therefore, the reduction in root colonization by AMF under water stress conditions can be attributed to a decrease in activity, spore germination, and mycelial development [73]. In addition, Sanaullah et al. [74] found that under water deficit conditions, the solubility of nutrients around the rhizosphere, the availability of which determines the activity of microorganisms such as AMF, decreases.
Oxidized monoterpenes and sesquiterpenes dominated the EO of sage with and without AMF. These groups are described as the main classes of EOs from sage. In AMF-inoculated plants, a decrease in oxidized monoterpenes was observed with an increase in oxidized sesquiterpenes at all irrigation levels. These results highlight the significant effect of AMF on the chemical profile of sage EO. Similarly, Tarraf et al. [18] stated that the chemical composition of the EO was drastically altered by Septoglomus viscosum, in which manool was the main component (28.13%), while α-thujone decreased (13.09%). These results suggest that AM symbiosis is a good candidate for promoting plant growth and essential oil composition. Khaliq et al. [23] explained that AMF symbiosis with thyme roots increases the area of mineral uptake as a result of extensive underground.
The main components recognized in the EO of sage with and without AMF were 1,8-cineole, α-thujone, β-thujone, camphor, E-caryophyllene, and viridiflorol. Colonization of roots with mycelium resulted in a significant increase in only 1,8-cineole and E-caryophyllene. Mycelium treatments had an overall positive effect on the accumulation of 1,8-cineole and E-caryophyllene in all levels of soil water content. In contrast, the fungi inoculation effect was positive for camphor only under water stress conditions (25% FC). It was clearly evident that an increase in irrigation rates in mycorrhizal plants reduced the amount of α-thujone and β-thujone. From an industrial perspective, essential oil with a lower thujone content is more desirable in the food sector due to potential health risks for consumers [18]. In this work, the thujone contents of the EOs tested are in accordance with ISO [75] (18.0–43.0 for α-thujone; 3.0 8.5% for β-thujone). Our study suggests that in the production of sage for the pharmaceutical industry from mycorrhizal and irrigated plants (optimally 75% FC), it is possible to obtain a high biomass yield and, at the same time, high-quality EO with low thujone content.

4.5. Correlations Between Parameters

In the present study, in the mycorrhiza of sage with AMF, the amount of fresh and air-dry biomass yield was mainly related to plant height and the number of leaves. A positive correlation (r = 0.59) was found between the percentage of EO in leaves and the estimated EO yield and the number of leaves per plant. The increase in plant shoot biomass in response to AM is widely recognized, and terpenoid accumulation in plants with AM is often associated with increased leaf biomass. Covello et al. [76] showed that terpenoids are synthesized and stored in the secretory structure of the leaf in trichomes. Studies have shown a significant correlation between trichome density and terpenoid accumulation in plants [5].

5. Conclusions

Herbaceous plant yields tend to be lower under water-deficient conditions. Our research indicates that irrigation can provide high yields of fresh and air-dry biomass and yields of sage leaves with higher essential oil content. Our conclusions regarding the response of sage plants to mycorrhizae are based on the range of irrigation levels tested. Most EO were found in the herb at moderate as well as high soil water levels (50%, 75% and 95% FC), while in the leaves, they were at 75% FC. The application of mycorrhiza increased air-dry biomass yield and leaf yield at each irrigation level. Mycorrhizal inoculation of sage roots at the 50% FC irrigation level was the most effective, as there was the most EO in the herb and leaves. AMF contributed to the higher content of sesquiterpenes in EO, but not monoterpenes. At a soil moisture level of 75% FC, both with and without AMF, there was a higher proportion of 1,8-cineole and camphor and a lower proportion of α-thujone and β-thujone. Root inoculation with AMF at an irrigation level of 75% FC reduced the proportion of α-thujone and β-thujone in EO.
Our study suggests that it is possible to obtain a high biomass yield and, at the same time, high-quality EO with low thujone content when sage is produced from mycorrhizal and irrigated plants (optimally 75% FC). Since the symbiotic effects of mycorrhiza are highly species-specific, it is necessary to direct future research to understanding plant metabolism in response to the cooperation of different AM species. A possible direction for further research could be a quantitative analysis of root colonization by AMF as a result of different irrigation levels, and to trace the interaction between AMF, plants, and soil microorganisms. By focusing future research on the aforementioned knowledge gaps in this area, mycorrhizal technology could open up new opportunities in sustainable medicinal plant growing practices, including water savings. In conclusion, AMFs are a powerful microbial resource that has great potential to improve the accumulation of medically important metabolites in host plants.

Author Contributions

Conceptualization, A.S. and R.N.-W.; methodology, A.S. and R.N.-W.; software, A.S.; validation, R.N.-W.; formal analysis, A.S., R.N.-W. and H.M.-R.; investigation, A.S. and R.N.-W.; resources, A.S. and H.M.-R.; data curation, H.M.-R. and S.I.-A.; writing—original draft preparation, A.S. and R.N.-W.; writing—review and editing, A.S., R.N.-W. and A.K.; visualization, H.M.-R., J.M.-G. and S.I.-A.; supervision, R.N.-W., A.K., S.I.-A. and A.F.; project administration, A.S., R.N.-W., J.M.-G. and A.F.; funding acquisition, R.N.-W. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Data Availability Statement

Data confirming the presented results are available from the authors and can be made available by contacting the corresponding author.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. The interaction effect of irrigation regimes and AMF inoculation on the number of sage leaves. Different letters above the columns denote significant differences among means (p < 0.05). Arbuscular mycorrhiza fungi (AMF); non-treated plants (non-AMF).
Figure 1. The interaction effect of irrigation regimes and AMF inoculation on the number of sage leaves. Different letters above the columns denote significant differences among means (p < 0.05). Arbuscular mycorrhiza fungi (AMF); non-treated plants (non-AMF).
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Figure 2. The interaction effect of irrigation regimes and AMF inoculation on the total air-dry herb (A) and total air-dry leaves (B) of sage. Different letters above the columns denote significant differences among means (p < 0.05). Arbuscular mycorrhiza fungi (AMF); non-treated plants (non-AMF).
Figure 2. The interaction effect of irrigation regimes and AMF inoculation on the total air-dry herb (A) and total air-dry leaves (B) of sage. Different letters above the columns denote significant differences among means (p < 0.05). Arbuscular mycorrhiza fungi (AMF); non-treated plants (non-AMF).
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Figure 3. The interaction effect of irrigation regimes and AMF inoculation on the EO in herbs (A) and EO in the leaves (B) of sage. Different letters above the columns denote significant differences among means (p < 0.05). Arbuscular mycorrhiza fungi (AMF); non-treated plants (non-AMF).
Figure 3. The interaction effect of irrigation regimes and AMF inoculation on the EO in herbs (A) and EO in the leaves (B) of sage. Different letters above the columns denote significant differences among means (p < 0.05). Arbuscular mycorrhiza fungi (AMF); non-treated plants (non-AMF).
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Figure 4. The interaction effect of irrigation regimes and AMF inoculation on the EO yield (A) and E-caryophyllene content (B). Different letters above the columns denote significant differences among means (p < 0.05). Arbuscular mycorrhiza fungi (AMF); non-treated plants (non-AMF).
Figure 4. The interaction effect of irrigation regimes and AMF inoculation on the EO yield (A) and E-caryophyllene content (B). Different letters above the columns denote significant differences among means (p < 0.05). Arbuscular mycorrhiza fungi (AMF); non-treated plants (non-AMF).
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Figure 5. The interaction effect of irrigation regimes and AMF inoculation on the 1,8-cineole (A) and camphor (B) content in EO. Different letters above the columns denote significant differences among means (p < 0.05). Arbuscular mycorrhiza fungi (AMF); non-treated plants (non-AMF).
Figure 5. The interaction effect of irrigation regimes and AMF inoculation on the 1,8-cineole (A) and camphor (B) content in EO. Different letters above the columns denote significant differences among means (p < 0.05). Arbuscular mycorrhiza fungi (AMF); non-treated plants (non-AMF).
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Figure 6. The interaction effect of irrigation regimes and AMF inoculation on α- thujone (A) and β-thujone (B) content in EO. Different letters above the columns denote significant differences among means (p < 0.05). Arbuscular mycorrhiza fungi (AMF); non-treated plants (non-AMF).
Figure 6. The interaction effect of irrigation regimes and AMF inoculation on α- thujone (A) and β-thujone (B) content in EO. Different letters above the columns denote significant differences among means (p < 0.05). Arbuscular mycorrhiza fungi (AMF); non-treated plants (non-AMF).
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Figure 7. Simple correlation coefficients between growth and yield attributes and EO content and composition.
Figure 7. Simple correlation coefficients between growth and yield attributes and EO content and composition.
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Table 1. Effect of irrigation regime and AMF inoculation on some morphological features of sage plants.
Table 1. Effect of irrigation regime and AMF inoculation on some morphological features of sage plants.
TreatmentsPlant HeightNumber
of Leaves		Number
of Branches		Total Biomass
Herb Yield		Total Leaves
Biomass
cm(Per Plant)(Per Plant)kg ha−1kg ha−1
Irrigation (I) (% FC)
95 ± 533.20 ± 3.61 a22.85 ± 0.60 a7.00 ± 0.70 a3273.70 ± 734.05 a2043.71 ± 225.11 a
75 ± 532.50 ± 1.71 a22.20 ± 3.28 a6.00 ± 0.88 b3188.40 ± 188.95 a2097.67 ± 124.06 a
50 ± 527.40 ± 1.34 b10.91 ± 3.39 b5.40 ± 0.34 c2655.60 ± 165.24 b1329.12 ± 112.56 b
25 ± 527.20 ± 1.98 b10.38 ± 2.41 b5.20 ± 0.98 c2686.60 ± 168.53 b1297.10 ± 122.98 b
Arbuscular mycorrhizal fungi (AMF)
AMF31.00 ± 4.37 a22.53 ± 3.29 a1.83 ± 0.83 a3036.35 ± 438 a1781.03 ± 446.11 a
non-AMF29.15 ± 2.39 a10.65 ± 5.32 b2.35 ± 0.35 a2865.80 ± 259 a1602.77 ± 363.87 a
Source of variation
I**********
AMFns***nsnsns
Interactionns**nsnsns
Explanations: ns (statistically insignificant differences); *, **, ***, respectively, significant at p < 0.05, p < 0.01, and p < 0.001. Arbuscular mycorrhiza fungi (AMF) treatment; non-treated plants (non-AMF). Values in columns followed by different letters are significantly different. Means ± standard deviation.
Table 2. Effect of irrigation regime and AMF inoculation on the sage yield and essential oil content.
Table 2. Effect of irrigation regime and AMF inoculation on the sage yield and essential oil content.
TreatmentsTotal of
Air-Dry Biomass		Total of
Air-Dry Leaves		EO Content
in Herb		EO Content
in Leaves		EO Yield
kg ha−1kg ha−1%%kg ha−1
Irrigation (I) (%FC)
95 ± 5460.22 ± 51 a320.13 ± 45 a0.778 ± 0.08 a0.894 ± 0.09 c18.27 ± 5.69 b
75 ± 5441.01 ± 42 a293.98 ± 24 a0.862 ± 0.12 a1.361 ± 0.17 a28.55 ± 6.89 a
50 ± 5257.99 ± 37 b129.72 ± 22 b0.771 ± 0.44 a1.132 ± 0.60 b15.05 ± 6.88 c
25 ± 5233.65 ± 59 b112.94 ± 24 b0.585 ± 0.32 b0.985 ± 0.12 c12.78 ± 3.85 d
Arbuscular mycorrhizal fungi (AMF)
AMF397.35 ± 55 a250.54 ± 89 a0.950 ± 0.16 a1.306 ± 0.33 a23.26 ± 5.69 a
non-AMF299.04 ± 64 b177.84 ± 72 b0.548 ± 0.25 b0.879 ± 0.24 b14.09 ± 6.89 b
Source of variation
I*************
AMF************
Interaction************
Explanations: *, **, *** are, respectively, significant at p < 0.05, p < 0.01, and p < 0.001. Arbuscular mycorrhiza fungi (AMF) treatment; non-treated plants (non-AMF). Values in the columns followed by different letters are significantly different. Means ± standard deviation.
Table 3. Compound groups of EO (percentages; mean ± SD) of S. officinalis after AMF treatment and non-AMF.
Table 3. Compound groups of EO (percentages; mean ± SD) of S. officinalis after AMF treatment and non-AMF.
Compound Groups Irrigation (%FC)
95 ± 575 ± 550 ± 525 ± 5
MH *non-AMF **10.69 ± 1.22 b z9.86 ± 1.23 b10.25 ± 1.25 b14.35 ± 1.67 a
AMF7.81 ± 0.74 b8.15 ± 1.01 b7.98 ± 0.80 b12.24 ± 1.67 a
OMnon-AMF56.44 ± 5.92 a58.19 ± 5.89 a52.98 ± 6.13 b49.44 ± 3.15 b
AMF38.15 ± 2.16 d48.59 ± 3.14 c32.98 ± 2.17 d33.34 ± 2.78 d
SHnon-AMF19.43 ± 1.56 b18.44 ± 1.22 c18.06 ± 1.56 c17.33 ± 1.15 c
AMF24.35 ± 2.13 a27.28 ± 1.89 a22.16± 2.03 b21.56 ± 2.67 b
OSnon-AMF9.65 ± 0.95 c7.74 ± 0.78 bc11.54 ± 0.89 b10.99 ± 0.75 b
AMF28.55 ± 1.55 a13.34 ± 0.59 b32.87 ± 1.33 a29.8 ± 1.45 a
ODnon-AMF1.70 ± 0.02 c2.70 ± 0.46 b2.10 ± 0.33 b4.20 ± 0.41 a
AMF0.12 ± 0.00 d1.09 ± 0.20 c2.45 ± 0.06 b2.26 ± 0.08 b
NCnon-AMF0.91 ± 0.01 a0.74 ± 0.07 a0.73 ± 0.01 a0.50 ± 0.03 b
AMF0.30 ± 0.00 a0.95 ± 0.09 a0.76 ± 0.01 a0.57± 0.06 a
Explanations: * MH—monoterpene hydrocarbons; OM—oxygenated monoterpenes; SH—sesquiterpene hydrocarbons; OS—oxygenated sesquiterpenes; OD—oxygenated diterpenes; NC—not identified compounds; ** non-treated plants (non-AMF); arbuscular mycorrhiza fungi (AMF); z—values in rows for particular compounds followed by different letters are significantly different. Means ± standard deviation.
Table 4. Effect of irrigation regime and AM inoculation on the chemical constituents of EO (%).
Table 4. Effect of irrigation regime and AM inoculation on the chemical constituents of EO (%).
Treatments1,8-Cineoleα-Thujoneβ-ThujoneCamphorE-CaryophylleneViridiflorol
Irrigation (I) (%FC)
95 ± 56.457 ± 0.68 b19.992 ± 2.07 ab7.800 ± 1.10 ab15.014 ± 1.81 b7.294 ± 1.46 b7.679 ± 1.12 b
75 ± 57.948 ± 0.67 a16.300 ± 4.37 c7.445 ± 1.33 c15.518 ± 2.71 a6.793 ± 1.85 c5.789 ± 0.97 c
50 ± 56.866 ± 0.81 b17.897 ± 4.36 b7.734 ± 1.34 b14.534 ± 2.52 b7.742 ± 1.11 a9.040 ± 1.11 a
25 ± 56.562 ± 0.72 b21.229 ± 4.16 a8.077 ± 1.56 a13.265 ± 2.50 c7.659 ± 0.96 a7.481 ± 1.11 b
Arbuscular mycorrhiza fungi (AMF)
AMF7.338 ± 1.27 a18.841 ± 5.93 a7.783 ± 1.04 a14.664 ± 3.99 a8.485 ± 1.27 a7.700 ± 1.15 a
non-AMF6.580 ± 1.17 b18.872 ± 5.54 a7.743 ± 1.34 a14.557 ± 2.63 a6.260 ± 1.33 b7.290 ± 1.25 a
Source of variation
I*****************
AMF***nsnsns***ns
Interaction**********ns
ns, *, **, *** are, respectively, not significant, significant at p < 0.05, p < 0.01, and p < 0.001. Arbuscular mycorrhiza fungi (AMF) treatment; non-treated plants (non-AMF). Values in columns followed by different letters are significantly different. Means ± standard deviation.
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Sałata, A.; Nurzyńska-Wierdak, R.; Kalisz, A.; Ibánez-Asensio, S.; Moreno-Ramón, H.; Majkowska-Gadomska, J.; Francke, A. Arbuscular Mycorrhizal Fungi (AMF) Influence Yield and Essential Oil Content and Composition of Sage (Salvia officinalis L.) Under Different Water Regimes. Agronomy 2025, 15, 2753. https://doi.org/10.3390/agronomy15122753

AMA Style

Sałata A, Nurzyńska-Wierdak R, Kalisz A, Ibánez-Asensio S, Moreno-Ramón H, Majkowska-Gadomska J, Francke A. Arbuscular Mycorrhizal Fungi (AMF) Influence Yield and Essential Oil Content and Composition of Sage (Salvia officinalis L.) Under Different Water Regimes. Agronomy. 2025; 15(12):2753. https://doi.org/10.3390/agronomy15122753

Chicago/Turabian Style

Sałata, Andrzej, Renata Nurzyńska-Wierdak, Andrzej Kalisz, Sara Ibánez-Asensio, Héctor Moreno-Ramón, Joanna Majkowska-Gadomska, and Anna Francke. 2025. "Arbuscular Mycorrhizal Fungi (AMF) Influence Yield and Essential Oil Content and Composition of Sage (Salvia officinalis L.) Under Different Water Regimes" Agronomy 15, no. 12: 2753. https://doi.org/10.3390/agronomy15122753

APA Style

Sałata, A., Nurzyńska-Wierdak, R., Kalisz, A., Ibánez-Asensio, S., Moreno-Ramón, H., Majkowska-Gadomska, J., & Francke, A. (2025). Arbuscular Mycorrhizal Fungi (AMF) Influence Yield and Essential Oil Content and Composition of Sage (Salvia officinalis L.) Under Different Water Regimes. Agronomy, 15(12), 2753. https://doi.org/10.3390/agronomy15122753

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