Co-application of arbuscular mycorrhizae fungi and putrescine improves essential oil production and drought tolerance cumin (Cuminum cyminum L.)

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Hamid Reza Anjam, Moslem Abdipour, Mehdi Hosseinifarahi This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4627617/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 27 Apr, 2026 Read the published version in Scientific Reports → Version 1 posted 12 You are reading this latest preprint version Abstract The effects of arbuscular mycorrhizae fungi (AMF), putrescine (Put), and their combination on essential oil (EO) content, chlorophyll (Chl), carotenoid and EO chemical compounds of cumin ( Cuminum cyminum L.) were studied under optimal and drought stress conditions. The foliar application of Put or inoculation with AMF significantly ( P ˂ 0.05) increased seed yield, EO content, EO yield, Chl a, Chl b, total Chl, and carotenoid. Moreover, AMF or Put application increased cuminaldehyde, limone, and β-Pinene and decreased γ-Terpinene, o-Cymene, and linalool concentration in cumin plants compared to non-treated plants in both optimal and drought stress conditions. However, the highest EO content, EO yield, Chl a, Chl b, total Chl, carotenoid, and major EO chemical components were obtained when AMF and Put were used together under optimal and drought stress conditions. This study showed that Put and AMF can be co-applied to the cumin plant to significantly improve drought tolerance in field conditions greatly . Biological sciences/Physiology Biological sciences/Plant sciences Chemical compounds Cumin Drought stress Mycorrhizae Putrescine Figures Figure 1 1. Introduction Cumin ( Cuminum cyminum L.), is an essential spice and medicinal plant from the Apiaceae family with a wide range of uses in various industries 1 . This plant is widely cultivated in arid and semi-arid regions due to its short growing season and low water requirement 2 . However, cumin plants face a variety of abiotic stresses during growth in these regions, such as drought stress, which can greatly affect their morphological and physiological characteristics and the concentration of their secondary metabolites and essential oil 3 . Limited water resources, besides progressive demand from food, pharmaceutical, and cosmetic industries, have encouraged researchers to improve water use in medicinal and aromatic plants with an emphasis on sustainable agriculture. It has been documented that defense systems in plants such as antioxidant may be insufficient to reduce the negative consequence of radical oxygen species (ROS) in stressful conditions such as drought stress 4 . Most recently, researchers have been drawn towards the application of biofertilizers to cope with drought stress. Biofertilizers are an emerging alternative considered a safer, cost-effective, and eco-friendly source of sustainable crop production with reduced chemical input 5 . Arbuscular mycorrhizal fungi (AMF) are known as the most important biofertilizers to improve the quantity and quality of plants by increasing the availability and maintaining the ecological balance of soil nutrients 6 . AMF can also efficiently increase plant tolerance to different stressful conditions, e.g., drought stress 7,8 , salinity 9 and heavy metals 10 , by set up a common symbiotic connection with roots 11 . Increased tolerance to drought stress in plants inoculated with AMF can be due to regulation of leaf stomatal conductivity, increase hydraulic conductivity of roots, reinstate ionic homeostasis, and balance biological process 12,13 . AMF alters the biosynthesis of essential oils by inducing the synthesis of secondary metabolites through the so-called induced systemic resistance (ISR) mechanism 14 . Polyamines (PAs) are a class of biogenic amines with multiple vital roles in the growth of plants. Three ubiquitous amines, viz., spermidine, spermine, and putrescine, are the most frequent PAs in plants and are widely used as plant growth regulators (PGRs) 15 . These osmotically active substances have been determined as potential candidates to ameliorate the adverse effects of environmental stresses 16 . The accumulation of PAs in plants exposed to environmental stresses and its vital role in stress tolerance have motivated researchers to apply PAs exogenously to cope with the adverse effects of various environmental stresses, such as drought 17 . Several reports have confirmed the role of PAs as a regulatory factor in improving the drought tolerance in medicinal plants, such as chamomile, thyme, fennel, and flame lily 17–20 . So far, there is no report on the effects of co-application of PAs and AMF on EO content and EO components in cumin plants under drought stress. Therefore, the main object of this research was to confirm whether AMF and Put as PBs can increase seed yield, EO content, and EO yield in Cuminum cyminum L. under normal irrigation and drought stress conditions. Furthermore, changing the EO profiles of Cuminum cyminum L. with the use of AMF and Put was investigated. 2. Materials and Methods 2.1. Study site and experimental layout A two-year field experiment was conducted at the Chamkhani Research Station, Yasouj, Iran (51°31ʹ06ʺ N, 30°41ʹ59ʺ E, Elevation 1740 m), with an average temperature of 16°C and 11.5°C and humidity of 47.6% and 49% during the growing season in 2018 and 2019, respectively. The soil at the experimental site was characterized as a clay loam with a hygroscopicity index of 7.3, a C:N of < 17, organic matter content of 0.96%, N content of 1%, available P content of 24 ppm, available K content of 337 ppm, and pH of 7.1. Planting was done by hand in individual plots (4m × 3m) with an inter-row spacing of 20 cm and a plant distance of 5 cm. Cumin plants were sown on 10 March 2018 and 13 March 2019 and harvested on 10 June 2018 and 15 June 2019. This research was performed in a factorial experiment according to a randomized complete block design (RCBD) with three replications. The factors were ( 1 ) irrigation treatment, including irrigation after 20% (normal irrigation) and 75% (drought stress) of the available soil water was depleted, ( 2 ) AMF inoculum ( Glomus intraradices ), including control (non-inoculation) and inoculation with the fungi, and ( 3 ) the foliar application of Put at three levels, i.e., control (0 mg L − 1 ), 1000, and 2000 mg L − 1 . All treatments were irrigated with a drip irrigation system every three days for five weeks to facilitate seedling establishment. Irrigation treatments were applied from April 16 and 19 in the first and second year, respectively, and continued throughout the growing season (end of June in both years). The water requirement (m 3 ) for each treatment was determined based on the soil water content (θi) using the TDR method (Time Domain Reflectometry, Model 4593) according to the following Eq. 2 1 : Vw: (θF.C − θi) × Bd×D ×A where Vw is the irrigation water volume, θF.C is the volume of soil moisture at field capacity (FC), θi is the soil water content, Bd is the soil bulk deity (kg/m 3 ), D is the rooting depth (0.3 m), and A is the main plot area (m 2 ). The AMF inoculation was down according to Khalediyan, et al. 22 with an inoculum of 35 g containing approximately 1450 spores. The AM fungi ( G. intraradices ) were obtained from the Soil and Water Research Institute, Iran. The foliar application of Put was performed twice. The first application was performed one week before drought treatment (April 9 and 12), and the second one was performed two weeks after the first application. Put was sprayed by an atomizer as the leaves were completely covered. 2.3. Plant sampling and data collecting At the flowering stage (May 10 and 13 in 2018 and 2019, respectively), 10–12 plants were randomly selected and the contents of chlorophyll (a, b, and total) and carotenoid pigments were measured. Chlorophyll a , b , and a + b (total) was measured based on 23 procedure. To measure the amount of chlorophyll, 1 g of leaf sample of each plot was beaten with 5 mL acetone 80% in a porcelain mortar. After purifying the resulting extract, the remaining leftovers were completely washed with 10 mL of acetone and passed through filter paper. Finally, the samples were as completely homogenized liquid to 10 mL volume. Optical absorbance was read by a spectrophotometer device at 663 and 645 (for chlorophyll a and b , respectively). Carotenoids content was determined using the method of Gu, et al. 24 . The seed yield was measured on plants harvested from 2 m 2 of each plot. 2.4. EO extraction and Gas chromatography/mass spectrometry (GC-MS) analysis EO was extracted using distillation with a glass Clevenger-type apparatus as recommended by the British Pharmacopoeia 25 . EO components analyses were down according to de Lira, et al. 26 with a Hewlett-Packard 6890 gas chromatograph. More details can be found in Sharafabad, et al. 27 . The retention indices (RI) were calculated using the standard solution of C8–C20 n-alkane (Fluka® Analytical, Munich, Germany) in hexane according to Adams 28 method. The bioactive constituents of cumin were confirmed by comparison of their RI with those reported in similar studies 29,30 . 2.5. Statistical analysis The data after testing for normality and homogeneity of variance, were subjected to a combined analysis of variance (ANOVA). The differences among the means were tested by Duncan’s multiple range test. Due to the insignificant ( P ≤ 0.05) effect of year on the traits, the average of the two-year data was used for further analyses. Statistical Analysis System (SAS) package version 9.4 31 was used for statistical analysis. 3. Results and Discussion 3.1. Seed yield The combined ANOVA indicated that the effects of water deficit treatments, putrescine (Put), arbuscular mycorrhizal fungi (AMF), and their interactions were significant on the seed yield (Table 1 ). However, the effect of year and it’s interaction with water deficit treatments, Put, and AMF were not significant ( p ≤ 0.05) on this trait. Duncan’s test indicated that the drought stress (25% FC) drastically reduced the seed yield of the cumin plants by 79.1% compared to the control (Table 2 ). The results are in line with the studies of Alipour, et al. 32 on fennel and the research of Jahani, et al. 33 on peppermint. Water deficit conditions, especially in the reproductive stage, result in the closure of stomatal apparatuses, which reduces chlorophyll florescence and then photosynthesis duration 34 . Such reduction in the photosynthesis period leads to a decrease in material mobilization to the seeds and the share of remobilization of material stored in the stem to seeds and a decrease in the seed yield 35,36 . Table 1 Combined analysis of variance related to the studied traits in two years SOV df Seed yield Essential oil (%) Essential oil yield Chlorophyll a Chlorophyll b Total chlorophyll Carotenoids Year 1 20480 2.731 45.43 0.119 0.112 0.255 0.155 Block (Year) 4 2876 0.421 11.59 0.070 0.034 0.038 0.207 Water deficit 1 49603** 2.221** 59.47** 0.881** 0.527** 1.427** 2.124** Year × Water deficit 1 2435 0.292 10.25 0.077 0.070 0.399 0.284 Block (Year × Water deficit) 4 493.2 0.084 2.487 0.032 0.017 0.058 0.053 Putrescin 2 7360** 1.212** 9.660** 0.454** 0.059** 0.408** 0.418** Mycorrhizal fungi 1 11139** 2.057** 13.92** 0.392** 0.103** 0.731** 0.399** Year × Putrescin 2 3202 0.354 4.131 0.139 0.027 0.080 0.098 Year × Mycorrhizal fungi 1 4116 0.511 2.820 0.087 0.049* 0.163 0.170 Water deficit × Putrescin 2 10152** 1.893** 15.86** 0.309** 0.098** 0.489** 0.513** Water deficit × Mycorrhizal fungi 1 17919** 2.416** 16.51** 0.508** 0.177** 0.574** 0.646** Putrescin × Mycorrhizal fungi 2 129159** 1.415** 17.16** 0.491** 0.057* 0.605** 0.558* Year × Water deficit × Putrescin 2 1239 0.456 5.101 0.162 0.021 0.161 0.138 Year × Water deficit × Mycorrhizal fungi 1 1134 0.197 2.487 0.053 0.038 0.257 0.164 Year × Putrescin × Mycorrhizal fungi 2 1722 0.260 7.125* 0.101 0.032 0.089 0.115 Water deficit × Putrescin × Mycorrhizal fungi 2 14297** 0.944** 12.80** 0.591** 0.135** 0.396** 0.453** Year × Water deficit × Putrescin × Mycorrhizal fungi 2 3202 0.396 3.230 0.169 0.031 0.203 0.140 Error 40 1050 0.158 1.673 0.053 0.011 0.065 0.046 CV† (%) 10.61 15.95 16.07 12.72 17.21 10.64 4.89 *Significant at the level of 0.05; **Significant at the level of 0.01; †CV = Coefficient of variance Table 2 Effects of polyamine and arbuscular mycorrhiza fungi (AMF) on traits in cumin plant under water deficit. Water deficit Putrescine Mycorrhizal fungi Seed yield (kg. ha − 1 ) Essential oil (%) Essential oil yield (kg. ha − 1 ) Chlorophyll a (mg g − 1 .FW) Chlorophyll b (mg g − 1 .FW) Total chlorophyll (mg g − 1 .FW) Carotenoids (mg g − 1 .FW) 80% available soil water 0 mg L − 1 Non-inoculation 308.89 de 1.91 h 5.89 f 1.44 e 0.41 c 1.85 gh 4.25 f AMF-inoculation 402.68 b 2.43 e 9.78 b 2.01 c 0.54 b 2.55 cd 4.8 d 1000 mg L − 1 Non-inoculation 322.06 d 2.01 g 6.47 e 1.52 e 0.45 bc 1.97 g 4.31 f AMF-inoculation 417.03 a 2.79 c 11.63 a 2.36 b 0.66 ab 3.02 b 5.79 a 2000 mg L − 1 Non-inoculation 374.37 c 2.17 f 8.12 c 1.91 c 0.47 bc 2.38 e 4.7 d AMF-inoculation 423.68 a 3.05 b 12.92 a 2.77 a 0.75 a 3.52 a 5.57 ab 30% available soil water 0 mg L − 1 Non-inoculation 172.44 i 2.13 f 3.67 i 0.96 fg 0.31 d 1.27 k 3.24 ij AMF-inoculation 267.61 g 2.61 d 6.98 d 1.87 d 0.39 c 2.26 f 3.68 h 1000 mg L − 1 Non-inoculation 178.35 i 2.16 f 3.85 h 1.12 f 0.36 cd 1.48 j 3.38 i AMF-inoculation 285.55 f 2.94 b 8.39 c 2.06 c 0.43 c 2.49 d 4.58 e 2000 mg L − 1 Non-inoculation 220.34 h 2.38 e 5.24 g 1.26 f 0.38 cd 1.64 i 3.94 g AMF-inoculation 291.1 f 3.31 a 9.63 b 2.15 bc 0.56 b 2.71 c 5.08 c Values marked with the same alphabets are not significantly different (LSD, p ≤ 0.05). Under non-stress conditions, the application of Put (1000 and 2000 mgL − 1 ) or AMF significantly increased the seed yield by 4.26, 21.19, and 30.36% compared to the non-treated plants, respectively (Table 2 ). A 3.43, 27.78 and 55.19% increase was recorded in the cumin plants treated with 1000 and 2000 mgL − 1 or inoculated with AMF over the non-treated plants under water deficit conditions, respectively. These results are consistent with the researchers who found a significant increase in seed yield when plants were inoculated with AMF under non-water stress conditions 37–39 and water deficit stress conditions 40,41 . The higher resistance of mycorrhizal plants to environmental stress can be related to the fact that AMF inoculation improves root architecture and the change of endogenous phytohormone contents in plants 42 . The improvement of root architecture that leads to the absorption of more water and nutrients from the soil is mainly attributed to the increase in the lateral root number, total root length, root surface area, and root volume in AMF-inoculated plants 43 . Compared to non-stress conditions, the cumin plants treated with Put and/or AMF under water deficit stress produced a higher seed yield. For example, Put application to the cumin plants in water stress conditions led to a 15.6% increase in this trait, while it led to a 12.73% increase in the seed yield in non-stress conditions (Table 2 ). Similarly, AMF under water stress conditions led to a significant increase compared to the non-stress conditions (55.19˃30.36%). Although foliar application of Put and AMF inoculation increased seed yield, it seems that cumin plants inoculated with AMF compared those treated with Put showed a greater extent of seed yield (on average, 42.77% compared to 14.16%). Interestingly, the Put and AMF treatments had a synergistic effect, where the highest seed yield was recorded in the plants treated with 2000 mgL − 1 Put and AMF. Numerous studies have been reported that the co-application of the two PBs can increase seed yield more than their individual application 44,45 . 3.2. Percentage of essential oil According to the results of combined ANOVA, the effects of water deficit stress, Put, AMF, and their interactions were significant on essential oil percentage (EO) ( p ˃0.01, Table 1 ). However, the effect of year and its interactions with other treatments was not significant on this trait ( p ˃0.05, Table 1 ). Interestingly, a significant increase (about 12%) in EO percentage with water deficit stress was observed compared with the plants under non-stress conditions (Table 2 ). The accumulation of secondary metabolites due to adverse environmental conditions, like drought, is a defense response to keep up cell balance and reduce the damaging results of stress 46 . In line with the results of the present study, in their research on the effect of drought stress on the EO percentage of Pelargonium graveolens , Mentha piperita L., and Melissa officinalis L., respectively, Amiri, et al. 47 , Jahani, et al. 33 , and Eshaghi Gorgi, et al. 7 reported that the EO percentage significantly increased with enhancing the intensity of drought stress. In addition, the results of this study showed that the application of Put increased the percentage of EO by 9.42 and 6.57% under non-stress and stress conditions, respectively. These results are consistent with those of Abd Elbar, et al. 48 , according to which the Thymus vulgaris L. plants treated with Put showed higher EO content under both non-drought and drought stress than the plants without Put application. Our findings showed a significant difference in response to Put concentrations (1000 and 2000 mgL − 1 ). The cumin plants treated with 2000 mgL − 1 Put had a higher percentage of EO than those treated with 1000 mgL − 1 of Put. Our results support the results of Mohammadi, et al. 18 , who obtained the highest increase in the percentage of EO under both non-drought stress and stress when the Thymus vulgaris L. plants were treated with the highest concentration of Put. Furthermore, the results showed that AMF colonization under non-stress and water deficit stress conditions caused a statistically significant increase of 27.23% and 20.83% in the EO percentage compared with the control treatment (Table 2 ). It has been documented that colonization with mycorrhizae can improve the EO percentage by improving soil microbial activity, increasing P availability, and producing plant growth regulators 22,49 . Under both non-stress and water deficit stress conditions, the cumin plants inoculated with AMF had a higher EO percentage than those treated with Put. However, the maximum rise in the EO percentage was exhibited by the co-application of AMF and Put compared to their individual use. 3.3. Essential oil yield The EO yield was remarkably ( p ˃0.01) impressed by the trial factors and their interactions (except year, Table 1 ). Duncan’s multiple range test showed that water deficit stress significantly reduced the EO yield by 60.99% in the cumin plants compared to the control. Basically, the EO yield in medicinal plants depends on the percentage of EO and the seed yield/flower yield, so changes in either factor can affect the EO yield 50 . According to the results, although the oil content increased in drought stress conditions, the seed yield decreased drastically (79.1%), leading to a decrease in the EO yield under drought stress conditions. Our results agree with those formerly reported by Mehrabi, et al. 51 on cumin and Alipour, et al. 32 on fennel, who reported that the EO yield was significantly influenced by drought stress. A pronounced increase was observed in the cumin plants inoculated with AMF under non-stress and water deficit stress conditions (66.04% and 90.19%, respectively, Table 2 ). The symbiosis between AMF and plants enhances phosphorous, iron, calcium, sulfur, copper, and zinc uptake and can elevate resistance to biotic and abiotic stresses 52 . The application of AMF, especially under drought stress, as an approach to increasing the EO yield has been reported in medicinal plants, such as Pelargonium graveolens L. 47 , Ocimum basilicum , and Satureja hortensis 22 , Foeniculum vulgare Mill 32 , and Melissa officinalis L. 7 . The cumin plants treated with Put also exhibited a significant increase compared to control under both non-stress and water deficit stress conditions. Owing to its tissue and outer layer stabilizing properties, its acid-neutralizing and antioxidant functions, Put can maintain a high water status, especially in dry environments, and can alleviate the impact on water status 53 . The pivotal role of Put in maintaining higher water status under drought stress and enhancing seed yield and EO yield have been reported by Mohammadi, et al. 18 , Zeynali, et al. 54 on Salvia officinalis L., and Nazarli and Naderi Arefi 55 on Matricaria chamomilla L. However, the maximum increase in EO yield was manifested by the cumin plants treated with both Put and AMF (Table 2 ). 3.4. Leaf chlorophyll content and carotenoid The results of combined ANOVA showed that the effects of water deficit stress, Put, AMF, and their interactions were significant on chlorophyll a , b , total chlorophyll, and carotenoid (Table 1 ). The cumin plants subjected to drought stress exhibited a notable reduction in above traits (Table 2 ). Chlorophyll a, b, total chlorophyll, and carotenoid of the cumin plants dropped significantly by 50%, 32.25%, 45.67%, and 31.17%, respectively, under induced water stress than under the non-stressed control plants (Table 2 ). Like our results, Mohammadi, et al. 18 found that the thyme plants exhibited 38.82, 51.11, and 41.61% lower chlorophyll a, chlorophyll b, and total chlorophyll contents under drought stress over the control, respectively. Chlorophyll, a key pigment to regulate the strength of the photosynthesis operation and crop production, is very sensitive to drought stress 56 . Drought stress can reduce chlorophyll content through destruction of chlorophyll substrate, chloroplast peroxidation, chlorophyll photo-oxidation, imbalance of protein complexes, inhibition of chlorophyll biosynthesis, and the increase in chlorophyllase activity Taïbi, et al. 57 . In this work, chlorophyll a was found to be more sensitive to drought stress than chlorophyll b (Table 2 ). Our results are in agreement with Kamrava, et al. 58 and Alipour, et al. 32 , who reported that chlorophyll a was more sensitive to water deficit and drastically decreased compared to chlorophyll b in rapeseed and fennel. The decline in carotenoid content due to drought stress has been reported in Melissa officinalis L. and Nigella sativa L. 7,59 . The malfunction of beta-carotene and the shaping of xanthine in the xanthophyll cycle has been reported as one of the reasons for carotenoid content reduction under drought stress conditions 59 . However, the cumin plants significantly benefitted from AMF inoculation, leading to increased chlorophyll a, b, total chlorophyll, and carotenoid compared to the control plants without AMF inoculation. For example, the inoculation of AMF led to an increase of 39.58%, 31.71%, 37.83%, and 12.94% under non-stress conditions and led to an increase of 94.79%, 25.81%, 77.95%, and 13.58% under water deficit stress conditions in chlorophyll a, b, total chlorophyll, and carotenoid, respectively (Table 2 ). Our results are in agreement with those of Zhu, et al. 60 and Hashem, et al. 61 , who reported that AMF inoculation increased chlorophyll content. Improving chlorophyll content by AMF inoculation, especially under drought stress, can be related to enhancing magnesium uptake and, consequently, improving plant growth 62 . The role of AMF in restoring the contents of pigments under drought stress, especially carotenoids, can be associated with the reduction of oxidative stress and ROS levels, thereby improving plant adaptation to drought stress 62 . Likewise, the cumin plants treated with Put showed an increase in chlorophyll a, b, total chlorophyll, and carotenoid by 19.09%, 12.19%, 17.56%, and 6% under non-stress conditions and an increase of 23.95%, 19.35%, 22.83%, and 12.96% under water deficit stress conditions, respectively. Similar to our results, Talaat, et al. 63 and Mohammadi, et al. 18 reported that the application of Put on Catharanthus roseus and Thymus vulgaris L. increased chlorophyll a, b, and carotenoid contents. Serafini-Fracassini, et al. 64 attributed the reduction of the destructive effects of drought stress due to spermine treatment in Lactuca sativa L. to an increase in TGase activity. The interactive effects of AMF and Put in both conditions were significant on chlorophyll a, b, total chlorophyll, and carotenoid ( p ˃0.05, Table 1 ). The highest chlorophyll a, b, total chlorophyll, and carotenoid contents were obtained when AMF and Put were applied together (Table 2 ). 3.5. Correlation coefficient The results of Pearson correlation between traits are shown as a heatmap in Fig. 1 . A strong positive correlation was found between the seed yield and EO yield (r 0.01 =0.883), chlorophyll a (r 0.01= 0.834), chlorophyll b (r 0.01 =0.851), total chlorophyll (r 0.01= 0.850), and carotenoids (r 0.01= 0.886). Therefore, an increase in seed yield will be associated with an increase in EO yield and vice versa. Furthermore, the positive and significant correlation between the seed yield and pigments showed the key role of these pigments, especially under drought stress, in gaining an acceptable seed yield. The importance of chlorophyll contents (a,b, and total chlorophyll) and carotenoids in resistance to environmental stress through eliminating free oxygen radicals 65 is consistent with the results of this study. 3.6. EO components The cumin EO samples in different treatments were subjected to a comparative chemical analysis through GC-MS. Cuminaldehyde was identified as the major compound, followed by limonene, γ- Terpinene, o-Cymene, linalool, and β-Pinene accounting for more than 45% of the total components (Table 3 ). In line with the results of the present study, Ebrahimabadi, et al. 29 and Ouryemchi, et al. 66 found that cuminaldehyde was the predominant component of cumin EO. However, the content of cuminaldehyde significantly varied under different experimental conditions. Table 3 Effects of polyamine and arbuscular mycorrhiza fungi (AMF) on traits in cumin plant under water deficit. Water deficit Putrescine Mycorrhizal fungi cuminaldehyde limonene γ- Terpinene o-Cymene linalool β-Pinene 80% available soil water 0 mg L − 1 Non-inoculation 14.25 e 12.08 f 11.1 f 6.74 e 5.84 d 4.41 g AMF-inoculation 16.33 c 13.82 b 12.25 d 7.94 ab 6.47 c 5.57 b 1000 mg L − 1 Non-inoculation 15.74 cd 13.37 c 10.87 fg 7.26 c 5.52 e 5.24 c AMF-inoculation 18.01 a 14.1a b 13.9 a 7.72 b 7.29 a 5.94 a 2000 mg L − 1 Non-inoculation 17.78 a 13.59 bc 13.01 b 6.92 d 6.77 b 4.97 d AMF-inoculation 17.59 ab 14.41 a 13.72 a 8.21 a 6.92 b 5.73 a 30% available soil water 0 mg L − 1 Non-inoculation 14.7 de 12.75 e 9.07 h 5.87 f 4.68 h 4.64 ef AMF-inoculation 16.06 c 13.45 c 11.36 e 7.21 c 5.84 d 4.96 d 1000 mg L − 1 Non-inoculation 15.31 d 13.05 d 10.41 g 7.01 cd 5.01 fg 4.81 e AMF-inoculation 16.88 bc 13.6 bc 12.43 c 7.52 bc 5.92 d 5.12 c 2000 mg L − 1 Non-inoculation 15.21 d 13.53 bc 11.32 e 7.61 b 5.21 f 5.06 cd AMF-inoculation 17.24 b 13.69 b 11.29 e 7.2 c 6.11 cd 5.22 c Values marked with the same alphabets are not significantly different (LSD, p ≤ 0.05). A significant variation in response to water deficit stress was observed among the major constituents of cumin EO. Cuminaldehyde, limonene, and β-Pinene increased by 3.15%, 5.54%, and 5.22% under water deficit stress conditions compared to non-stress conditions, while γ- Terpinene, o-Cymene, and linalool decreased by 22.38%, 14.82%, and 24.78%, respectively (Table 3 ). These results are consistent with Rebey, et al. 67 , who found a notable increase in cuminaldehyde, limonene, and β-Pinene under water deficit stress in cumin plants. Mohammadi, et al. 18 reported similar results for Thymus vulgaris L. under drought stress, according to which the plants exhibited lower rates of γ- Terpinene, p -Cymene, and linalool compared with well-watered plants. Importantly, cuminaldehyde, limonene, β-Pinene, γ- Terpinene, o-Cymene, and linalool in cumin plants treated with AMF increased by 14.59%, 14.40%, 10.36%, 17.80%, 10.78%, and 26.30% under non-stress conditions and by 9.25%, 5.49%, 25.24%, 22.83%, 24.78% and 6.89% under water deficit stress conditions compared to the control. In agreement with our results, Ebrahimabadi, et al. 29 showed that AMF application significantly increased cuminaldehyde, limonene, β-Pinene, γ- Terpinene, o-Cymene, and linalool in cumin plants. Changes in the ratio of EO compounds with AMF inoculation under normal and drought stress conditions have been reported for black cumin 68 , lemon balm 7 , and Ajowan 40 . Likewise, the foliar application of Put under both non-stress and stressful conditions led to an average increase of 7.31%, 6.52%, 8.42%, 13.58%, 0.78%, and 11.24% in cuminaldehyde, limonene, β-Pinene, γ- Terpinene, o-Cymene, and linalool, respectively (Table 3 ). Similar observations were reported by Mohammadi, et al. 18 , who showed a higher accumulation of some major EO constituents in thyme under non-drought stress and drought stress conditions due to the treatment with Put compared to the control. Changes in the proportion of EO compounds with the use of different concentrations of Put have been reported for lavender 27 , marigold 69 , and sage 54 . However, the maximum increase in cuminaldehyde, limonene, β-Pinene, γ- Terpinene, o-Cymene, and linalool was exhibited by the co-application of AMF and Put (2000 mgL − 1 ). 4. Conclusion Our results provide insights into understanding the effects of AMF and Put on seed yield, EO yield, EO content, and composition in cumin under drought stress. Based on the results, AMF and Put can be considered advantageous for enhancing cumin production in arid and semi-arid regions because they improve photosynthetic pigments, seed yield, and EO yield accumulation. Conclusively, due to the synergistic effects of AMF and Put, the co-application of AMF and Put in cumin plants is suggested to improve plant tolerance to drought stress and enhance commercial value. Abbreviations AMF, Arbuscular mycorrhizae fungi; Put, putrescine; EO, Essential oil; Chl, chlorophyll, ROS, Radical oxygen species; ISR, Induced systemic resistance; PGR's, Plant growth regulators, PAs, Polyamines; PBs, Plantbiostimulants; ANOVA, Analysis of variance; RCBD, Randomized complete block design; TDR, Time Domain Reflectometry; F.C, Field capacity; GC-MS, Gas chromatography-mass spectrometry; RI, Retention index; SAS, Statistical analysis system. Declarations Declaration of Competing Interest The authors declare that they have no conflict of interest. Author Contribution Hamid Reza Anjam: Investigation, data collection, and original draft preparation. Moslem Abdipour & Mehdi Hosseinifarahi: Conceptualization, Methodology, Validation, Investigation, Supervision, Software, Formal analysis, Writing - review & editing. Data Availability We confirm that we provide same data availability statement in the manuscript and in submission system. The datasets generated and/or analysed during the current study are available in the [data] Supplementary material, [https://submission.springernature.com/submission/54fab868-4b2f-4982-930a-2fdc2d92e63c/file/121cc2f9-feca-465c-95c4-760ed58d87ea] References Lodha, S. & Mawar, R. Cumin wilt management: a review. J. Spice. Aromat. Crop 23, 145–155 (2014). Kumar, S. et al. Understanding Cuminum cyminum: An important seed spice crop of arid and semi-arid regions. Int. J. Seed Spices 5, 1–19 (2015). Hajmohammadnia Ghalibaf, K. et al. Effects of Planting Date and Deficit Irrigation on Water Use Efficiency of Cumin (Cuminum cyminum L.) at two different Densities in Mashhad Conditions. 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Communications in Soil Science and Plant Analysis 55, 343–364 (2024). Additional Declarations No competing interests reported. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-4627617","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":327716959,"identity":"8eec3576-3623-4627-8432-100448a8a1f2","order_by":0,"name":"Hamid Reza Anjam","email":"","orcid":"","institution":"Islamic Azad University","correspondingAuthor":false,"prefix":"","firstName":"Hamid","middleName":"Reza","lastName":"Anjam","suffix":""},{"id":327716962,"identity":"8f3f4331-429c-49ac-af74-d765d767020e","order_by":1,"name":"Moslem Abdipour","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA30lEQVRIiWNgGAWjYHACNgYeAxDNfAAhlkCcFjYkdYS1gGmIRsJAt4H52YM3BXfkDG6f+fi5osJGjoH98AOGh3twazE7wGZuOMfgmbHBudzNkmfOpBkz8KQZMCQ8w6eFwUyax+Bw4oYzvBskG9sOJzYw5AD9cgCfFvZvUC08j382/vuf2MD/hpAWHpgtPGySjQ0HEhskCNlymKcc7BfJM2xmlg3Hko3ZJJ4ZHMCr5Xj7tgdv/tyR4zvD/PhmQ42dHD9/8sOHP/BoYWAGk0gq2FC5OAExakbBKBgFo2DEAgD1KFGWYHz3MQAAAABJRU5ErkJggg==","orcid":"","institution":"Agricultural Research, Education and Extension Organization (AREEO)","correspondingAuthor":true,"prefix":"","firstName":"Moslem","middleName":"","lastName":"Abdipour","suffix":""},{"id":327716963,"identity":"3c1236d0-0f6f-4939-a309-820f3981ea9c","order_by":2,"name":"Mehdi Hosseinifarahi","email":"","orcid":"","institution":"Islamic Azad University","correspondingAuthor":false,"prefix":"","firstName":"Mehdi","middleName":"","lastName":"Hosseinifarahi","suffix":""}],"badges":[],"createdAt":"2024-06-24 05:30:33","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4627617/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4627617/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1038/s41598-026-43180-w","type":"published","date":"2026-04-27T15:57:54+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":60696642,"identity":"4ea3fdb8-2453-466a-8ba8-864cd4cf6a6c","added_by":"auto","created_at":"2024-07-19 16:34:47","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":90174,"visible":true,"origin":"","legend":"\u003cp\u003eHeat-map matrix of the correlation between traits of cumin from different treatments. Each square indicates \u003cem\u003er\u003c/em\u003e (Pearson’s correlation coefficient of a pair of traits).\u003c/p\u003e","description":"","filename":"OnlineFig.1.png","url":"https://assets-eu.researchsquare.com/files/rs-4627617/v1/f1c3c20a2d42fbf9e6439f8b.png"},{"id":108437810,"identity":"645b709e-ecfb-46a1-9625-b88fe0d8fcff","added_by":"auto","created_at":"2026-05-04 16:03:44","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":629787,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4627617/v1/6e08056e-8d66-4ec4-95ce-eadd07860e60.pdf"},{"id":60696643,"identity":"613cba17-a5dd-4bdf-8675-5bc40c7f81e9","added_by":"auto","created_at":"2024-07-19 16:34:47","extension":"xlsx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":20073,"visible":true,"origin":"","legend":"","description":"","filename":"Data.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-4627617/v1/7d62b4a26c8d62b7824c3ae3.xlsx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Co-application of arbuscular mycorrhizae fungi and putrescine improves essential oil production and drought tolerance cumin (Cuminum cyminum L.)","fulltext":[{"header":"1. Introduction","content":"\u003cp\u003eCumin (\u003cem\u003eCuminum cyminum\u003c/em\u003e L.), is an essential spice and medicinal plant from the Apiaceae family with a wide range of uses in various industries \u003csup\u003e1\u003c/sup\u003e. This plant is widely cultivated in arid and semi-arid regions due to its short growing season and low water requirement \u003csup\u003e2\u003c/sup\u003e. However, cumin plants face a variety of abiotic stresses during growth in these regions, such as drought stress, which can greatly affect their morphological and physiological characteristics and the concentration of their secondary metabolites and essential oil \u003csup\u003e3\u003c/sup\u003e. Limited water resources, besides progressive demand from food, pharmaceutical, and cosmetic industries, have encouraged researchers to improve water use in medicinal and aromatic plants with an emphasis on sustainable agriculture.\u003c/p\u003e \u003cp\u003eIt has been documented that defense systems in plants such as antioxidant may be insufficient to reduce the negative consequence of radical oxygen species (ROS) in stressful conditions such as drought stress \u003csup\u003e4\u003c/sup\u003e. Most recently, researchers have been drawn towards the application of biofertilizers to cope with drought stress. Biofertilizers are an emerging alternative considered a safer, cost-effective, and eco-friendly source of sustainable crop production with reduced chemical input \u003csup\u003e5\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eArbuscular mycorrhizal fungi (AMF) are known as the most important biofertilizers to improve the quantity and quality of plants by increasing the availability and maintaining the ecological balance of soil nutrients \u003csup\u003e6\u003c/sup\u003e. AMF can also efficiently increase plant tolerance to different stressful conditions, e.g., drought stress \u003csup\u003e7,8\u003c/sup\u003e, salinity \u003csup\u003e9\u003c/sup\u003e and heavy metals \u003csup\u003e10\u003c/sup\u003e, by set up a common symbiotic connection with roots \u003csup\u003e11\u003c/sup\u003e. Increased tolerance to drought stress in plants inoculated with AMF can be due to regulation of leaf stomatal conductivity, increase hydraulic conductivity of roots, reinstate ionic homeostasis, and balance biological process \u003csup\u003e12,13\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eAMF alters the biosynthesis of essential oils by inducing the synthesis of secondary metabolites through the so-called induced systemic resistance (ISR) mechanism \u003csup\u003e14\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003ePolyamines (PAs) are a class of biogenic amines with multiple vital roles in the growth of plants. Three ubiquitous amines, viz., spermidine, spermine, and putrescine, are the most frequent PAs in plants and are widely used as plant growth regulators (PGRs) \u003csup\u003e15\u003c/sup\u003e. These osmotically active substances have been determined as potential candidates to ameliorate the adverse effects of environmental stresses \u003csup\u003e16\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eThe accumulation of PAs in plants exposed to environmental stresses and its vital role in stress tolerance have motivated researchers to apply PAs exogenously to cope with the adverse effects of various environmental stresses, such as drought \u003csup\u003e17\u003c/sup\u003e. Several reports have confirmed the role of PAs as a regulatory factor in improving the drought tolerance in medicinal plants, such as chamomile, thyme, fennel, and flame lily \u003csup\u003e17\u0026ndash;20\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eSo far, there is no report on the effects of co-application of PAs and AMF on EO content and EO components in cumin plants under drought stress. Therefore, the main object of this research was to confirm whether AMF and Put as PBs can increase seed yield, EO content, and EO yield in \u003cem\u003eCuminum cyminum\u003c/em\u003e L. under normal irrigation and drought stress conditions. Furthermore, changing the EO profiles of \u003cem\u003eCuminum cyminum\u003c/em\u003e L. with the use of AMF and Put was investigated.\u003c/p\u003e"},{"header":"2. Materials and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003e2.1. Study site and experimental layout\u003c/h2\u003e \u003cp\u003eA two-year field experiment was conducted at the Chamkhani Research Station, Yasouj, Iran (51\u0026deg;31ʹ06ʺ N, 30\u0026deg;41ʹ59ʺ E, Elevation 1740 m), with an average temperature of 16\u0026deg;C and 11.5\u0026deg;C and humidity of 47.6% and 49% during the growing season in 2018 and 2019, respectively. The soil at the experimental site was characterized as a clay loam with a hygroscopicity index of 7.3, a C:N of \u0026lt;\u0026thinsp;17, organic matter content of 0.96%, N content of 1%, available P content of 24 ppm, available K content of 337 ppm, and pH of 7.1.\u003c/p\u003e \u003cp\u003ePlanting was done by hand in individual plots (4m \u0026times; 3m) with an inter-row spacing of 20 cm and a plant distance of 5 cm. Cumin plants were sown on 10 March 2018 and 13 March 2019 and harvested on 10 June 2018 and 15 June 2019.\u003c/p\u003e \u003cp\u003eThis research was performed in a factorial experiment according to a randomized complete block design (RCBD) with three replications. The factors were (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e) irrigation treatment, including irrigation after 20% (normal irrigation) and 75% (drought stress) of the available soil water was depleted, (\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e) AMF inoculum (\u003cem\u003eGlomus intraradices\u003c/em\u003e), including control (non-inoculation) and inoculation with the fungi, and (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e) the foliar application of Put at three levels, i.e., control (0 mg L\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), 1000, and 2000 mg L\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eAll treatments were irrigated with a drip irrigation system every three days for five weeks to facilitate seedling establishment. Irrigation treatments were applied from April 16 and 19 in the first and second year, respectively, and continued throughout the growing season (end of June in both years).\u003c/p\u003e \u003cp\u003eThe water requirement (m\u003csup\u003e3\u003c/sup\u003e) for each treatment was determined based on the soil water content (θi) using the TDR method (Time Domain Reflectometry, Model 4593) according to the following Eq.\u0026nbsp;2\u003csup\u003e1\u003c/sup\u003e:\u003c/p\u003e \u003cp\u003eVw: (θF.C\u0026thinsp;\u0026minus;\u0026thinsp;θi) \u0026times; Bd\u0026times;D \u0026times;A\u003c/p\u003e \u003cp\u003ewhere \u003cem\u003eVw\u003c/em\u003e is the irrigation water volume, \u003cem\u003eθF.C\u003c/em\u003e is the volume of soil moisture at field capacity (FC), \u003cem\u003eθi\u003c/em\u003e is the soil water content, \u003cem\u003eBd\u003c/em\u003e is the soil bulk deity (kg/m\u003csup\u003e3\u003c/sup\u003e), \u003cem\u003eD\u003c/em\u003e is the rooting depth (0.3 m), and \u003cem\u003eA\u003c/em\u003e is the main plot area (m\u003csup\u003e2\u003c/sup\u003e).\u003c/p\u003e \u003cp\u003eThe AMF inoculation was down according to Khalediyan, et al. \u003csup\u003e22\u003c/sup\u003e with an inoculum of 35 g containing approximately 1450 spores. The AM fungi (\u003cem\u003eG. intraradices\u003c/em\u003e) were obtained from the Soil and Water Research Institute, Iran.\u003c/p\u003e \u003cp\u003eThe foliar application of Put was performed twice. The first application was performed one week before drought treatment (April 9 and 12), and the second one was performed two weeks after the first application. Put was sprayed by an atomizer as the leaves were completely covered.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003e2.3. Plant sampling and data collecting\u003c/h2\u003e \u003cp\u003eAt the flowering stage (May 10 and 13 in 2018 and 2019, respectively), 10\u0026ndash;12 plants were randomly selected and the contents of chlorophyll (a, b, and total) and carotenoid pigments were measured.\u003c/p\u003e \u003cp\u003eChlorophyll \u003cem\u003ea\u003c/em\u003e, \u003cem\u003eb\u003c/em\u003e, and a\u0026thinsp;+\u0026thinsp;b (total) was measured based on \u003csup\u003e23\u003c/sup\u003e procedure. To measure the amount of chlorophyll, 1 g of leaf sample of each plot was beaten with 5 mL acetone 80% in a porcelain mortar. After purifying the resulting extract, the remaining leftovers were completely washed with 10 mL of acetone and passed through filter paper. Finally, the samples were as completely homogenized liquid to 10 mL volume. Optical absorbance was read by a spectrophotometer device at 663 and 645 (for chlorophyll \u003cem\u003ea\u003c/em\u003e and \u003cem\u003eb\u003c/em\u003e, respectively). Carotenoids content was determined using the method of Gu, et al. \u003csup\u003e24\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eThe seed yield was measured on plants harvested from 2 m\u003csup\u003e2\u003c/sup\u003e of each plot.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003e2.4. EO extraction and Gas chromatography/mass spectrometry (GC-MS) analysis\u003c/h2\u003e \u003cp\u003eEO was extracted using distillation with a glass Clevenger-type apparatus as recommended by the British Pharmacopoeia \u003csup\u003e25\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eEO components analyses were down according to de Lira, et al. \u003csup\u003e26\u003c/sup\u003e with a Hewlett-Packard 6890 gas chromatograph. More details can be found in Sharafabad, et al. \u003csup\u003e27\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eThe retention indices (RI) were calculated using the standard solution of C8\u0026ndash;C20 n-alkane (Fluka\u0026reg; Analytical, Munich, Germany) in hexane according to Adams \u003csup\u003e28\u003c/sup\u003e method. The bioactive constituents of cumin were confirmed by comparison of their RI with those reported in similar studies \u003csup\u003e29,30\u003c/sup\u003e.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003e2.5. Statistical analysis\u003c/h2\u003e \u003cp\u003eThe data after testing for normality and homogeneity of variance, were subjected to a combined analysis of variance (ANOVA). The differences among the means were tested by Duncan\u0026rsquo;s multiple range test. Due to the insignificant (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026le;\u0026thinsp;0.05) effect of year on the traits, the average of the two-year data was used for further analyses. Statistical Analysis System (SAS) package version 9.4 \u003csup\u003e31\u003c/sup\u003e was used for statistical analysis.\u003c/p\u003e \u003c/div\u003e"},{"header":"3. Results and Discussion","content":"\u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003e3.1. Seed yield\u003c/h2\u003e \u003cp\u003eThe combined ANOVA indicated that the effects of water deficit treatments, putrescine (Put), arbuscular mycorrhizal fungi (AMF), and their interactions were significant on the seed yield (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). However, the effect of year and it\u0026rsquo;s interaction with water deficit treatments, Put, and AMF were not significant (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026le;\u0026thinsp;0.05) on this trait. Duncan\u0026rsquo;s test indicated that the drought stress (25% FC) drastically reduced the seed yield of the cumin plants by 79.1% compared to the control (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The results are in line with the studies of Alipour, et al. \u003csup\u003e32\u003c/sup\u003e on fennel and the research of Jahani, et al. \u003csup\u003e33\u003c/sup\u003e on peppermint. Water deficit conditions, especially in the reproductive stage, result in the closure of stomatal apparatuses, which reduces chlorophyll florescence and then photosynthesis duration \u003csup\u003e34\u003c/sup\u003e. Such reduction in the photosynthesis period leads to a decrease in material mobilization to the seeds and the share of remobilization of material stored in the stem to seeds and a decrease in the seed yield \u003csup\u003e35,36\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eCombined analysis of variance related to the studied traits in two years\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"9\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSOV\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003edf\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSeed yield\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eEssential oil (%)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eEssential oil yield\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eChlorophyll a\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eChlorophyll b\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eTotal chlorophyll\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eCarotenoids\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eYear\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e20480\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e2.731\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e45.43\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.119\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0.112\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0.255\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.155\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBlock (Year)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2876\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.421\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e11.59\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.070\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0.034\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0.038\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.207\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWater deficit\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e49603**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e2.221**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e59.47**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.881**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0.527**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e1.427**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e2.124**\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eYear \u0026times; Water deficit\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2435\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.292\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e10.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.077\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0.070\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0.399\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.284\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBlock (Year \u0026times; Water deficit)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e493.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.084\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2.487\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.032\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0.017\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0.058\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.053\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePutrescin\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e7360**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1.212**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e9.660**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.454**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0.059**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0.408**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.418**\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMycorrhizal fungi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e11139**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e2.057**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e13.92**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.392**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0.103**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0.731**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.399**\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eYear \u0026times; Putrescin\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3202\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.354\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e4.131\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.139\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0.027\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0.080\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.098\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eYear \u0026times; Mycorrhizal fungi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e4116\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.511\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2.820\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.087\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0.049*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0.163\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.170\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWater deficit \u0026times; Putrescin\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e10152**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1.893**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e15.86**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.309**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0.098**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0.489**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.513**\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWater deficit \u0026times; Mycorrhizal fungi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e17919**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e2.416**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e16.51**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.508**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0.177**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0.574**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.646**\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePutrescin \u0026times; Mycorrhizal fungi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e129159**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1.415**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e17.16**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.491**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0.057*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0.605**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.558*\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eYear \u0026times; Water deficit \u0026times; Putrescin\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1239\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.456\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e5.101\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.162\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0.021\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0.161\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.138\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eYear \u0026times; Water deficit \u0026times; Mycorrhizal fungi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1134\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.197\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2.487\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.053\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0.038\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0.257\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.164\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eYear \u0026times; Putrescin \u0026times; Mycorrhizal fungi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1722\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.260\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e7.125*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.101\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0.032\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0.089\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.115\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWater deficit \u0026times; Putrescin \u0026times; Mycorrhizal fungi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e14297**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.944**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e12.80**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.591**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0.135**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0.396**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.453**\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eYear \u0026times; Water deficit \u0026times; Putrescin \u0026times; Mycorrhizal fungi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3202\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.396\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3.230\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.169\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0.031\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0.203\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.140\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eError\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1050\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.158\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1.673\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.053\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0.011\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0.065\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.046\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCV\u0026dagger; (%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e10.61\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e15.95\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e16.07\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e12.72\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e17.21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e10.64\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e4.89\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"9\"\u003e*Significant at the level of 0.05; **Significant at the level of 0.01; \u0026dagger;CV\u0026thinsp;=\u0026thinsp;Coefficient of variance\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eEffects of polyamine and arbuscular mycorrhiza fungi (AMF) on traits in cumin plant under water deficit.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"10\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWater deficit\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePutrescine\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eMycorrhizal fungi\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSeed yield (kg. ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eEssential oil (%)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEssential oil yield (kg. ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eChlorophyll a (mg g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e.FW)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eChlorophyll b\u003c/p\u003e \u003cp\u003e(mg g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e.FW)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eTotal chlorophyll (mg g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e.FW)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e \u003cp\u003eCarotenoids (mg g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e.FW)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"5\" rowspan=\"6\"\u003e \u003cp\u003e80% available soil water\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e0 mg L\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNon-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e308.89\u003csup\u003ede\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.91\u003csup\u003eh\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e5.89\u003csup\u003ef\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1.44\u003csup\u003ee\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.41\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e1.85\u003csup\u003egh\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e4.25\u003csup\u003ef\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAMF-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e402.68\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.43\u003csup\u003ee\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e9.78\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2.01\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.54\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e2.55\u003csup\u003ecd\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e4.8\u003csup\u003ed\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e1000 mg L\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNon-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e322.06\u003csup\u003ed\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.01\u003csup\u003eg\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e6.47\u003csup\u003ee\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1.52\u003csup\u003ee\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.45\u003csup\u003ebc\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e1.97\u003csup\u003eg\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e4.31\u003csup\u003ef\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAMF-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e417.03\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.79\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e11.63\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2.36\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.66\u003csup\u003eab\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e3.02\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e5.79\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e2000 mg L\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNon-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e374.37\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.17\u003csup\u003ef\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e8.12\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1.91\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.47\u003csup\u003ebc\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e2.38\u003csup\u003ee\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e4.7\u003csup\u003ed\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAMF-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e423.68\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3.05\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e12.92\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2.77\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.75\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e3.52\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e5.57\u003csup\u003eab\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"5\" rowspan=\"6\"\u003e \u003cp\u003e30% available soil water\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e0 mg L\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNon-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e172.44\u003csup\u003ei\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.13\u003csup\u003ef\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e3.67\u003csup\u003ei\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.96\u003csup\u003efg\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.31\u003csup\u003ed\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e1.27\u003csup\u003ek\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e3.24\u003csup\u003eij\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAMF-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e267.61\u003csup\u003eg\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.61\u003csup\u003ed\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e6.98\u003csup\u003ed\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1.87\u003csup\u003ed\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.39\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e2.26\u003csup\u003ef\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e3.68\u003csup\u003eh\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e1000 mg L\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNon-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e178.35\u003csup\u003ei\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.16\u003csup\u003ef\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e3.85\u003csup\u003eh\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1.12\u003csup\u003ef\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.36\u003csup\u003ecd\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e1.48\u003csup\u003ej\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e3.38\u003csup\u003ei\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAMF-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e285.55\u003csup\u003ef\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.94\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e8.39\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2.06\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.43\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e2.49\u003csup\u003ed\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e4.58\u003csup\u003ee\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e2000 mg L\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNon-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e220.34\u003csup\u003eh\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.38\u003csup\u003ee\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e5.24\u003csup\u003eg\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1.26\u003csup\u003ef\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.38\u003csup\u003ecd\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e1.64\u003csup\u003ei\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e3.94\u003csup\u003eg\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAMF-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e291.1\u003csup\u003ef\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3.31\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e9.63\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2.15\u003csup\u003ebc\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.56\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e2.71\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e5.08\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"10\"\u003eValues marked with the same alphabets are not significantly different (LSD, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026le;\u0026thinsp;0.05).\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eUnder non-stress conditions, the application of Put (1000 and 2000 mgL\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) or AMF significantly increased the seed yield by 4.26, 21.19, and 30.36% compared to the non-treated plants, respectively (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). A 3.43, 27.78 and 55.19% increase was recorded in the cumin plants treated with 1000 and 2000 mgL\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e or inoculated with AMF over the non-treated plants under water deficit conditions, respectively. These results are consistent with the researchers who found a significant increase in seed yield when plants were inoculated with AMF under non-water stress conditions \u003csup\u003e37\u0026ndash;39\u003c/sup\u003e and water deficit stress conditions \u003csup\u003e40,41\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eThe higher resistance of mycorrhizal plants to environmental stress can be related to the fact that AMF inoculation improves root architecture and the change of endogenous phytohormone contents in plants \u003csup\u003e42\u003c/sup\u003e. The improvement of root architecture that leads to the absorption of more water and nutrients from the soil is mainly attributed to the increase in the lateral root number, total root length, root surface area, and root volume in AMF-inoculated plants \u003csup\u003e43\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eCompared to non-stress conditions, the cumin plants treated with Put and/or AMF under water deficit stress produced a higher seed yield. For example, Put application to the cumin plants in water stress conditions led to a 15.6% increase in this trait, while it led to a 12.73% increase in the seed yield in non-stress conditions (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Similarly, AMF under water stress conditions led to a significant increase compared to the non-stress conditions (55.19˃30.36%). Although foliar application of Put and AMF inoculation increased seed yield, it seems that cumin plants inoculated with AMF compared those treated with Put showed a greater extent of seed yield (on average, 42.77% compared to 14.16%). Interestingly, the Put and AMF treatments had a synergistic effect, where the highest seed yield was recorded in the plants treated with 2000 mgL\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e Put and AMF.\u003c/p\u003e \u003cp\u003eNumerous studies have been reported that the co-application of the two PBs can increase seed yield more than their individual application \u003csup\u003e44,45\u003c/sup\u003e.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec9\" class=\"Section2\"\u003e \u003ch2\u003e3.2. Percentage of essential oil\u003c/h2\u003e \u003cp\u003eAccording to the results of combined ANOVA, the effects of water deficit stress, Put, AMF, and their interactions were significant on essential oil percentage (EO) (\u003cem\u003ep\u003c/em\u003e˃0.01, Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). However, the effect of year and its interactions with other treatments was not significant on this trait (\u003cem\u003ep\u003c/em\u003e˃0.05, Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eInterestingly, a significant increase (about 12%) in EO percentage with water deficit stress was observed compared with the plants under non-stress conditions (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The accumulation of secondary metabolites due to adverse environmental conditions, like drought, is a defense response to keep up cell balance and reduce the damaging results of stress \u003csup\u003e46\u003c/sup\u003e. In line with the results of the present study, in their research on the effect of drought stress on the EO percentage of \u003cem\u003ePelargonium graveolens\u003c/em\u003e, \u003cem\u003eMentha piperita\u003c/em\u003e L., and \u003cem\u003eMelissa officinalis\u003c/em\u003e L., respectively, Amiri, et al. \u003csup\u003e47\u003c/sup\u003e, Jahani, et al. \u003csup\u003e33\u003c/sup\u003e, and Eshaghi Gorgi, et al. \u003csup\u003e7\u003c/sup\u003e reported that the EO percentage significantly increased with enhancing the intensity of drought stress.\u003c/p\u003e \u003cp\u003eIn addition, the results of this study showed that the application of Put increased the percentage of EO by 9.42 and 6.57% under non-stress and stress conditions, respectively. These results are consistent with those of Abd Elbar, et al. \u003csup\u003e48\u003c/sup\u003e, according to which the \u003cem\u003eThymus vulgaris\u003c/em\u003e L. plants treated with Put showed higher EO content under both non-drought and drought stress than the plants without Put application.\u003c/p\u003e \u003cp\u003eOur findings showed a significant difference in response to Put concentrations (1000 and 2000 mgL\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e). The cumin plants treated with 2000 mgL\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e Put had a higher percentage of EO than those treated with 1000 mgL\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e of Put. Our results support the results of Mohammadi, et al. \u003csup\u003e18\u003c/sup\u003e, who obtained the highest increase in the percentage of EO under both non-drought stress and stress when the \u003cem\u003eThymus vulgaris\u003c/em\u003e L. plants were treated with the highest concentration of Put.\u003c/p\u003e \u003cp\u003eFurthermore, the results showed that AMF colonization under non-stress and water deficit stress conditions caused a statistically significant increase of 27.23% and 20.83% in the EO percentage compared with the control treatment (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). It has been documented that colonization with mycorrhizae can improve the EO percentage by improving soil microbial activity, increasing P availability, and producing plant growth regulators \u003csup\u003e22,49\u003c/sup\u003e. Under both non-stress and water deficit stress conditions, the cumin plants inoculated with AMF had a higher EO percentage than those treated with Put. However, the maximum rise in the EO percentage was exhibited by the co-application of AMF and Put compared to their individual use.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec10\" class=\"Section2\"\u003e \u003ch2\u003e3.3. Essential oil yield\u003c/h2\u003e \u003cp\u003eThe EO yield was remarkably (\u003cem\u003ep\u003c/em\u003e˃0.01) impressed by the trial factors and their interactions (except year, Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Duncan\u0026rsquo;s multiple range test showed that water deficit stress significantly reduced the EO yield by 60.99% in the cumin plants compared to the control. Basically, the EO yield in medicinal plants depends on the percentage of EO and the seed yield/flower yield, so changes in either factor can affect the EO yield \u003csup\u003e50\u003c/sup\u003e. According to the results, although the oil content increased in drought stress conditions, the seed yield decreased drastically (79.1%), leading to a decrease in the EO yield under drought stress conditions. Our results agree with those formerly reported by Mehrabi, et al. \u003csup\u003e51\u003c/sup\u003e on cumin and Alipour, et al. \u003csup\u003e32\u003c/sup\u003e on fennel, who reported that the EO yield was significantly influenced by drought stress.\u003c/p\u003e \u003cp\u003eA pronounced increase was observed in the cumin plants inoculated with AMF under non-stress and water deficit stress conditions (66.04% and 90.19%, respectively, Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The symbiosis between AMF and plants enhances phosphorous, iron, calcium, sulfur, copper, and zinc uptake and can elevate resistance to biotic and abiotic stresses \u003csup\u003e52\u003c/sup\u003e. The application of AMF, especially under drought stress, as an approach to increasing the EO yield has been reported in medicinal plants, such as \u003cem\u003ePelargonium graveolens\u003c/em\u003e L. \u003csup\u003e47\u003c/sup\u003e, \u003cem\u003eOcimum basilicum\u003c/em\u003e, and \u003cem\u003eSatureja hortensis\u003c/em\u003e \u003csup\u003e22\u003c/sup\u003e, \u003cem\u003eFoeniculum vulgare\u003c/em\u003e Mill \u003csup\u003e32\u003c/sup\u003e, and \u003cem\u003eMelissa officinalis\u003c/em\u003e L. \u003csup\u003e7\u003c/sup\u003e. The cumin plants treated with Put also exhibited a significant increase compared to control under both non-stress and water deficit stress conditions.\u003c/p\u003e \u003cp\u003eOwing to its tissue and outer layer stabilizing properties, its acid-neutralizing and antioxidant functions, Put can maintain a high water status, especially in dry environments, and can alleviate the impact on water status \u003csup\u003e53\u003c/sup\u003e. The pivotal role of Put in maintaining higher water status under drought stress and enhancing seed yield and EO yield have been reported by Mohammadi, et al. \u003csup\u003e18\u003c/sup\u003e, Zeynali, et al. \u003csup\u003e54\u003c/sup\u003e on \u003cem\u003eSalvia officinalis\u003c/em\u003e L., and Nazarli and Naderi Arefi \u003csup\u003e55\u003c/sup\u003e on \u003cem\u003eMatricaria chamomilla\u003c/em\u003e L. However, the maximum increase in EO yield was manifested by the cumin plants treated with both Put and AMF (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003e3.4. Leaf chlorophyll content and carotenoid\u003c/h2\u003e \u003cp\u003eThe results of combined ANOVA showed that the effects of water deficit stress, Put, AMF, and their interactions were significant on chlorophyll \u003cem\u003ea\u003c/em\u003e, \u003cem\u003eb\u003c/em\u003e, total chlorophyll, and carotenoid (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The cumin plants subjected to drought stress exhibited a notable reduction in above traits (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Chlorophyll a, b, total chlorophyll, and carotenoid of the cumin plants dropped significantly by 50%, 32.25%, 45.67%, and 31.17%, respectively, under induced water stress than under the non-stressed control plants (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Like our results, Mohammadi, et al. \u003csup\u003e18\u003c/sup\u003e found that the thyme plants exhibited 38.82, 51.11, and 41.61% lower chlorophyll a, chlorophyll b, and total chlorophyll contents under drought stress over the control, respectively.\u003c/p\u003e \u003cp\u003eChlorophyll, a key pigment to regulate the strength of the photosynthesis operation and crop production, is very sensitive to drought stress \u003csup\u003e56\u003c/sup\u003e. Drought stress can reduce chlorophyll content through destruction of chlorophyll substrate, chloroplast peroxidation, chlorophyll photo-oxidation, imbalance of protein complexes, inhibition of chlorophyll biosynthesis, and the increase in chlorophyllase activity Ta\u0026iuml;bi, et al. \u003csup\u003e57\u003c/sup\u003e. In this work, chlorophyll a was found to be more sensitive to drought stress than chlorophyll b (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Our results are in agreement with Kamrava, et al. \u003csup\u003e58\u003c/sup\u003e and Alipour, et al. \u003csup\u003e32\u003c/sup\u003e, who reported that chlorophyll a was more sensitive to water deficit and drastically decreased compared to chlorophyll b in rapeseed and fennel.\u003c/p\u003e \u003cp\u003eThe decline in carotenoid content due to drought stress has been reported in \u003cem\u003eMelissa officinalis\u003c/em\u003e L. and \u003cem\u003eNigella sativa\u003c/em\u003e L. \u003csup\u003e7,59\u003c/sup\u003e. The malfunction of beta-carotene and the shaping of xanthine in the xanthophyll cycle has been reported as one of the reasons for carotenoid content reduction under drought stress conditions \u003csup\u003e59\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eHowever, the cumin plants significantly benefitted from AMF inoculation, leading to increased chlorophyll a, b, total chlorophyll, and carotenoid compared to the control plants without AMF inoculation. For example, the inoculation of AMF led to an increase of 39.58%, 31.71%, 37.83%, and 12.94% under non-stress conditions and led to an increase of 94.79%, 25.81%, 77.95%, and 13.58% under water deficit stress conditions in chlorophyll a, b, total chlorophyll, and carotenoid, respectively (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Our results are in agreement with those of Zhu, et al. \u003csup\u003e60\u003c/sup\u003e and Hashem, et al. \u003csup\u003e61\u003c/sup\u003e, who reported that AMF inoculation increased chlorophyll content. Improving chlorophyll content by AMF inoculation, especially under drought stress, can be related to enhancing magnesium uptake and, consequently, improving plant growth \u003csup\u003e62\u003c/sup\u003e. The role of AMF in restoring the contents of pigments under drought stress, especially carotenoids, can be associated with the reduction of oxidative stress and ROS levels, thereby improving plant adaptation to drought stress \u003csup\u003e62\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eLikewise, the cumin plants treated with Put showed an increase in chlorophyll a, b, total chlorophyll, and carotenoid by 19.09%, 12.19%, 17.56%, and 6% under non-stress conditions and an increase of 23.95%, 19.35%, 22.83%, and 12.96% under water deficit stress conditions, respectively.\u003c/p\u003e \u003cp\u003eSimilar to our results, Talaat, et al. \u003csup\u003e63\u003c/sup\u003e and Mohammadi, et al. \u003csup\u003e18\u003c/sup\u003e reported that the application of Put on \u003cem\u003eCatharanthus roseus\u003c/em\u003e and \u003cem\u003eThymus vulgaris\u003c/em\u003e L. increased chlorophyll a, b, and carotenoid contents. Serafini-Fracassini, et al. \u003csup\u003e64\u003c/sup\u003e attributed the reduction of the destructive effects of drought stress due to spermine treatment in \u003cem\u003eLactuca sativa\u003c/em\u003e L. to an increase in TGase activity. The interactive effects of AMF and Put in both conditions were significant on chlorophyll a, b, total chlorophyll, and carotenoid (\u003cem\u003ep\u003c/em\u003e˃0.05, Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The highest chlorophyll a, b, total chlorophyll, and carotenoid contents were obtained when AMF and Put were applied together (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003e3.5. Correlation coefficient\u003c/h2\u003e \u003cp\u003eThe results of Pearson correlation between traits are shown as a heatmap in Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e. A strong positive correlation was found between the seed yield and EO yield (r\u003csub\u003e0.01\u003c/sub\u003e=0.883), chlorophyll a (r\u003csub\u003e0.01=\u003c/sub\u003e0.834), chlorophyll b (r\u003csub\u003e0.01\u003c/sub\u003e=0.851), total chlorophyll (r\u003csub\u003e0.01=\u003c/sub\u003e0.850), and carotenoids (r\u003csub\u003e0.01=\u003c/sub\u003e0.886). Therefore, an increase in seed yield will be associated with an increase in EO yield and vice versa. Furthermore, the positive and significant correlation between the seed yield and pigments showed the key role of these pigments, especially under drought stress, in gaining an acceptable seed yield. The importance of chlorophyll contents (a,b, and total chlorophyll) and carotenoids in resistance to environmental stress through eliminating free oxygen radicals \u003csup\u003e65\u003c/sup\u003e is consistent with the results of this study.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003e3.6. EO components\u003c/h2\u003e \u003cp\u003eThe cumin EO samples in different treatments were subjected to a comparative chemical analysis through GC-MS. Cuminaldehyde was identified as the major compound, followed by limonene, γ- Terpinene, o-Cymene, linalool, and β-Pinene accounting for more than 45% of the total components (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). In line with the results of the present study, Ebrahimabadi, et al. \u003csup\u003e29\u003c/sup\u003e and Ouryemchi, et al. \u003csup\u003e66\u003c/sup\u003e found that cuminaldehyde was the predominant component of cumin EO. However, the content of cuminaldehyde significantly varied under different experimental conditions.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eEffects of polyamine and arbuscular mycorrhiza fungi (AMF) on traits in cumin plant under water deficit.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"9\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWater deficit\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePutrescine\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eMycorrhizal fungi\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003ecuminaldehyde\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003elimonene\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eγ- Terpinene\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eo-Cymene\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003elinalool\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eβ-Pinene\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"5\" rowspan=\"6\"\u003e \u003cp\u003e80% available soil water\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e0 mg L\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNon-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e14.25\u003csup\u003ee\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e12.08\u003csup\u003ef\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e11.1\u003csup\u003ef\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e6.74\u003csup\u003ee\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e5.84\u003csup\u003ed\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e4.41\u003csup\u003eg\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAMF-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e16.33\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e13.82\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e12.25\u003csup\u003ed\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e7.94\u003csup\u003eab\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e6.47\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e5.57\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e1000 mg L\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNon-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e15.74\u003csup\u003ecd\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e13.37\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e10.87\u003csup\u003efg\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e7.26\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e5.52\u003csup\u003ee\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e5.24\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAMF-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e18.01\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e14.1a\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e13.9\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e7.72\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e7.29\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e5.94\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e2000 mg L\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNon-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e17.78\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e13.59\u003csup\u003ebc\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e13.01\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e6.92\u003csup\u003ed\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e6.77\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e4.97\u003csup\u003ed\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAMF-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e17.59\u003csup\u003eab\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e14.41\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e13.72\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e8.21\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e6.92\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e5.73\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"5\" rowspan=\"6\"\u003e \u003cp\u003e30% available soil water\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e0 mg L\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNon-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e14.7\u003csup\u003ede\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e12.75\u003csup\u003ee\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e9.07\u003csup\u003eh\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e5.87\u003csup\u003ef\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e4.68\u003csup\u003eh\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e4.64\u003csup\u003eef\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAMF-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e16.06\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e13.45\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e11.36\u003csup\u003ee\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e7.21\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e5.84\u003csup\u003ed\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e4.96\u003csup\u003ed\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e1000 mg L\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNon-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e15.31\u003csup\u003ed\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e13.05\u003csup\u003ed\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e10.41\u003csup\u003eg\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e7.01\u003csup\u003ecd\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e5.01\u003csup\u003efg\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e4.81\u003csup\u003ee\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAMF-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e16.88\u003csup\u003ebc\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e13.6\u003csup\u003ebc\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e12.43\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e7.52\u003csup\u003ebc\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e5.92\u003csup\u003ed\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e5.12\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e2000 mg L\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNon-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e15.21\u003csup\u003ed\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e13.53\u003csup\u003ebc\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e11.32\u003csup\u003ee\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e7.61\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e5.21\u003csup\u003ef\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e5.06\u003csup\u003ecd\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAMF-inoculation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e17.24\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e13.69\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e11.29\u003csup\u003ee\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e7.2\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e6.11\u003csup\u003ecd\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e5.22\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"9\"\u003eValues marked with the same alphabets are not significantly different (LSD, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026le;\u0026thinsp;0.05).\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eA significant variation in response to water deficit stress was observed among the major constituents of cumin EO. Cuminaldehyde, limonene, and β-Pinene increased by 3.15%, 5.54%, and 5.22% under water deficit stress conditions compared to non-stress conditions, while γ- Terpinene, o-Cymene, and linalool decreased by 22.38%, 14.82%, and 24.78%, respectively (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThese results are consistent with Rebey, et al. \u003csup\u003e67\u003c/sup\u003e, who found a notable increase in cuminaldehyde, limonene, and β-Pinene under water deficit stress in cumin plants. Mohammadi, et al. \u003csup\u003e18\u003c/sup\u003e reported similar results for \u003cem\u003eThymus vulgaris\u003c/em\u003e L. under drought stress, according to which the plants exhibited lower rates of γ- Terpinene, \u003cem\u003ep\u003c/em\u003e-Cymene, and linalool compared with well-watered plants.\u003c/p\u003e \u003cp\u003eImportantly, cuminaldehyde, limonene, β-Pinene, γ- Terpinene, o-Cymene, and linalool in cumin plants treated with AMF increased by 14.59%, 14.40%, 10.36%, 17.80%, 10.78%, and 26.30% under non-stress conditions and by 9.25%, 5.49%, 25.24%, 22.83%, 24.78% and 6.89% under water deficit stress conditions compared to the control. In agreement with our results, Ebrahimabadi, et al. \u003csup\u003e29\u003c/sup\u003e showed that AMF application significantly increased cuminaldehyde, limonene, β-Pinene, γ- Terpinene, o-Cymene, and linalool in cumin plants. Changes in the ratio of EO compounds with AMF inoculation under normal and drought stress conditions have been reported for black cumin \u003csup\u003e68\u003c/sup\u003e, lemon balm \u003csup\u003e7\u003c/sup\u003e, and Ajowan \u003csup\u003e40\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eLikewise, the foliar application of Put under both non-stress and stressful conditions led to an average increase of 7.31%, 6.52%, 8.42%, 13.58%, 0.78%, and 11.24% in cuminaldehyde, limonene, β-Pinene, γ- Terpinene, o-Cymene, and linalool, respectively (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Similar observations were reported by Mohammadi, et al. \u003csup\u003e18\u003c/sup\u003e, who showed a higher accumulation of some major EO constituents in thyme under non-drought stress and drought stress conditions due to the treatment with Put compared to the control. Changes in the proportion of EO compounds with the use of different concentrations of Put have been reported for lavender \u003csup\u003e27\u003c/sup\u003e, marigold \u003csup\u003e69\u003c/sup\u003e, and sage \u003csup\u003e54\u003c/sup\u003e. However, the maximum increase in cuminaldehyde, limonene, β-Pinene, γ- Terpinene, o-Cymene, and linalool was exhibited by the co-application of AMF and Put (2000 mgL\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e).\u003c/p\u003e \u003c/div\u003e"},{"header":"4. Conclusion","content":"\u003cp\u003eOur results provide insights into understanding the effects of AMF and Put on seed yield, EO yield, EO content, and composition in cumin under drought stress. Based on the results, AMF and Put can be considered advantageous for enhancing cumin production in arid and semi-arid regions because they improve photosynthetic pigments, seed yield, and EO yield accumulation. Conclusively, due to the synergistic effects of AMF and Put, the co-application of AMF and Put in cumin plants is suggested to improve plant tolerance to drought stress and enhance commercial value.\u003c/p\u003e"},{"header":"Abbreviations","content":"\u003cp\u003e AMF, Arbuscular mycorrhizae fungi; Put, putrescine; EO, Essential oil; Chl, chlorophyll, ROS, Radical oxygen species; ISR, Induced systemic resistance; PGR's, Plant growth regulators, PAs, Polyamines; PBs, Plantbiostimulants; ANOVA, Analysis of variance; RCBD, Randomized complete block design; TDR, Time Domain Reflectometry; F.C, Field capacity; GC-MS, Gas chromatography-mass spectrometry; RI, Retention index; SAS, Statistical analysis system.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e \u003ch2\u003eDeclaration of Competing Interest\u003c/h2\u003e \u003cp\u003eThe authors declare that they have no conflict of interest.\u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eHamid Reza Anjam: Investigation, data collection, and original draft preparation. Moslem Abdipour \u0026amp; Mehdi Hosseinifarahi: Conceptualization, Methodology, Validation, Investigation, Supervision, Software, Formal analysis, Writing - review \u0026amp; editing.\u003c/p\u003e\u003ch2\u003eData Availability\u003c/h2\u003e\u003cp\u003eWe confirm that we provide same data availability statement in the manuscript and in submission system. The datasets generated and/or analysed during the current study are available in the [data] Supplementary material, [https://submission.springernature.com/submission/54fab868-4b2f-4982-930a-2fdc2d92e63c/file/121cc2f9-feca-465c-95c4-760ed58d87ea]\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eLodha, S. \u0026amp; Mawar, R. Cumin wilt management: a review. J. Spice. Aromat. 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Journal of Crop Breeding 9, 95\u0026ndash;104 (2017).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKabiri, R., Farahbakhsh, H. \u0026amp; Nasibi, F. Effect of drought stress on physiological and biochemical characteristics of Nigella sativa L. Iranian Journal of Medicinal and Aromatic Plants Research 30, 600\u0026ndash;610 (2014).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZhu, X. Q., Tang, M. \u0026amp; Zhang, H. Arbuscular mycorrhizal fungi enhanced the growth, photosynthesis, and calorific value of black locust under salt stress. Photosynthetica 55, 378\u0026ndash;385 (2017).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHashem, A. \u003cem\u003eet al.\u003c/em\u003e The interaction between arbuscular mycorrhizal fungi and endophytic bacteria enhances plant growth of Acacia gerrardii under salt stress. Frontiers in microbiology 7, 1089 (2016).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAl-Arjani, A.-B. F., Hashem, A. \u0026amp; Abd_Allah, E. F. Arbuscular mycorrhizal fungi modulates dynamics tolerance expression to mitigate drought stress in Ephedra foliata Boiss. Saudi Journal of Biological Sciences 27, 380\u0026ndash;394 (2020).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eTalaat, I. M., Bekheta, M. \u0026amp; Mahgoub, M. H. Physiological response of periwinkle plants (Catharanthus roseus L.) to tryptophan and putrescine. International Journal of Agriculture and Biology 7, 210\u0026ndash;213 (2005).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSerafini-Fracassini, D., Di Sandro, A. \u0026amp; Del Duca, S. Spermine delays leaf senescence in Lactuca sativa and prevents the decay of chloroplast photosystems. Plant Physiology and Biochemistry 48, 602\u0026ndash;611 (2010).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGill, S. S. \u0026amp; Tuteja, N. Reactive oxygen species and antioxidant machinery in abiotic stress tolerance in crop plants. Plant physiology and biochemistry 48, 909\u0026ndash;930 (2010).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eOuryemchi, I. \u003cem\u003eet al.\u003c/em\u003e GC-MS Characterization, Antioxidant, Antimicrobial and Insecticidal Potential of Moroccan Cuminum Cyminum L. Essential Oil. Tropical Journal of Natural Product Research 8 (2024).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eRebey, I. B. \u003cem\u003eet al.\u003c/em\u003e Effect of drought on the biochemical composition and antioxidant activities of cumin (Cuminum cyminum L.) seeds. Industrial Crops and Products 36, 238\u0026ndash;245 (2012).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDarakeh, S. A. S. S., Weisany, W., Tahir, N. A.-R. \u0026amp; Schenk, P. M. Physiological and biochemical responses of black cumin to vermicompost and plant biostimulants: Arbuscular mycorrhizal and plant growth-promoting rhizobacteria. Industrial Crops and Products 188, 115557 (2022).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBeheshti, F., Abdipour, M., Hosseinifarahi, M. \u0026amp; Kelidari, A. Characterization of Lavender Essential Oil and Its Chemical Composition Intercropped with Marigold as Affected by Polyamines and Arbuscular Mycorrhizal Fungi Inoculation. Communications in Soil Science and Plant Analysis 55, 343\u0026ndash;364 (2024).\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"scientific-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"scirep","sideBox":"Learn more about [Scientific Reports](http://www.nature.com/srep/)","snPcode":"","submissionUrl":"","title":"Scientific Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Scientific Reports","inReviewEnabled":true,"inReviewRevisionsEnabled":true},"keywords":"Chemical compounds, Cumin, Drought stress, Mycorrhizae, Putrescine","lastPublishedDoi":"10.21203/rs.3.rs-4627617/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4627617/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe effects of arbuscular mycorrhizae fungi (AMF), putrescine (Put), and their combination on essential oil (EO) content, chlorophyll (Chl), carotenoid and EO chemical compounds of cumin (\u003cem\u003eCuminum cyminum\u003c/em\u003e L.) were studied under optimal and drought stress conditions. The foliar application of Put or inoculation with AMF significantly (\u003cem\u003eP\u003c/em\u003e ˂ 0.05) increased seed yield, EO content, EO yield, Chl a, Chl b, total Chl, and carotenoid. Moreover, AMF or Put application increased cuminaldehyde, limone, and β-Pinene and decreased γ-Terpinene, o-Cymene, and linalool concentration in cumin plants compared to non-treated plants in both optimal and drought stress conditions. However, the highest EO content, EO yield, Chl a, Chl b, total Chl, carotenoid, and major EO chemical components were obtained when AMF and Put were used together under optimal and drought stress conditions. This study showed that Put and AMF can be co-applied to the cumin plant to significantly improve drought tolerance in field conditions greatly\u003ca class=\"FNLink\" href=\"#Fn1\" id=\"#FNLinkFn1\"\u003e\u003c/a\u003e.\u003c/p\u003e","manuscriptTitle":"Co-application of arbuscular mycorrhizae fungi and putrescine improves essential oil production and drought tolerance cumin (Cuminum cyminum L.)","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-07-19 16:34:42","doi":"10.21203/rs.3.rs-4627617/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2024-10-28T05:42:41+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2024-10-24T05:30:03+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2024-10-21T10:36:35+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"318062911178060866871322399775198691693","date":"2024-10-20T08:22:02+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2024-10-10T21:40:15+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"306841366843553074064198541524791719197","date":"2024-09-22T22:06:09+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"242218776100511100208054911544493085974","date":"2024-09-22T13:58:42+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2024-09-22T13:51:46+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2024-09-18T11:50:47+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"","date":"2024-06-27T16:22:07+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2024-06-26T08:54:15+00:00","index":"","fulltext":""},{"type":"submitted","content":"Scientific Reports","date":"2024-06-24T05:29:13+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"scientific-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"scirep","sideBox":"Learn more about [Scientific Reports](http://www.nature.com/srep/)","snPcode":"","submissionUrl":"","title":"Scientific Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Scientific Reports","inReviewEnabled":true,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"eebc2dd9-f5d2-45a7-a251-260a168d8a00","owner":[],"postedDate":"July 19th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[{"id":34685470,"name":"Biological sciences/Physiology"},{"id":34685471,"name":"Biological sciences/Plant sciences"}],"tags":[],"updatedAt":"2026-05-04T16:02:41+00:00","versionOfRecord":{"articleIdentity":"rs-4627617","link":"https://doi.org/10.1038/s41598-026-43180-w","journal":{"identity":"scientific-reports","isVorOnly":false,"title":"Scientific Reports"},"publishedOn":"2026-04-27 15:57:54","publishedOnDateReadable":"April 27th, 2026"},"versionCreatedAt":"2024-07-19 16:34:42","video":"","vorDoi":"10.1038/s41598-026-43180-w","vorDoiUrl":"https://doi.org/10.1038/s41598-026-43180-w","workflowStages":[]},"version":"v1","identity":"rs-4627617","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-4627617","identity":"rs-4627617","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}