Mycorrhizal fungal colonization promotes apparent growth and physiology of Alhagi sparsifolia seedlings under salt or drought stress at vulnerable developmental stage

Mycorrhizal fungal colonization promotes apparent growth and physiology of Alhagi sparsifolia seedlings under salt or drought stress at vulnerable developmental stage | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Mycorrhizal fungal colonization promotes apparent growth and physiology of Alhagi sparsifolia seedlings under salt or drought stress at vulnerable developmental stage Xiaonan Chen, Aili Yilinuer, Xiaodong Ma, Haiou Wang, Dawuti Maigepiretiguli This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-2374175/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 05 Apr, 2023 Read the published version in Plant Growth Regulation → Version 1 posted 6 You are reading this latest preprint version Abstract Alhagi sparsifolia is a dominant species in the lower Tarim River desert ecosystem and an important mycorrhizal symbiont. However, it is unclear how its seedlings cope with salt and drought stresses and develop effective survival strategies with the assistance of arbuscular mycorrhizal fungi (AMF) during the vulnerable growth stage. Therefore, in this study, we investigated the effects of salt or drought stress on the growth rate; nutrient uptake; photosynthesis; and the levels of antioxidant enzymes, osmoregulatory substances, and hormones in A. sparsifolia seedlings. We included following six groups: control/CK-NM (without salt or drought stress and no AMF inoculation), D-NM (drought stress and no AMF inoculation), S-NM (salt stress and no AMF inoculation), CK-AM (AMF inoculation and no salt or drought stress), D-AM (drought stress and AMF inoculation), and S-AM (salt stress and AMF inoculation). The results revealed that AMF inoculation promoted seedling growth, particularly root growth and phosphorus nutrient uptake, in A. sparsifolia . Salt and drought stresses negatively affected the growth, photosynthetic capacity, and nutrient accumulation in the above- and below-ground parts of the seedlings and stimulated the antioxidant defense system and accumulation of osmoregulatory substances in them. AMF inoculation under salt and drought stresses could alleviate toxic symptoms in A. sparsifolia by promoting root growth, enhancing nutrient uptake, activating antioxidant enzyme activity, and regulating hormonal levels. These effects of AMF were mainly reflected in root growth under drought stress and antioxidant enzyme activity under salt stress. The beneficial effect of AMF under salt stress was better than that under drought stress. This study demonstrated that AMF plays a significant role in assisting A. sparsifolia seedlings to quickly pass through the vulnerable growth stage under salt and drought stresses. Therefore, A. sparsifolia seedlings with AMF have potential application in restoration of desert ecosystem. The Tarim River Basin Alhagi sparsifolia seedlings Plant–Microbe Interaction Antioxidant enzyme system Phytohormones Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Introduction Salt and drought stresses are some of the most frequent and severe abiotic stresses in desert ecosystems. They negatively affect many physiological and biochemical indicators and growth and development in plants (Abderrahim et al. 2020 ). The lower Tarim River is a typical damaged desert ecosystem in the arid zone, and artificial water transfer projects have been implemented for 22 years to solve the problem of ecological water shortage. Downstream desert riparian forests are located in extremely arid inland areas, often with high salt accumulation in their artificially delivered surface and groundwater. This causes severe stress to the plant seedlings for their survival and growth (Zhang et al. 2022 ; Chen et al. 2021 ; Li et al. 2020 ). A. sparsifolia is a perennial herb in the legume family and a dominant herb in the riparian forests of the lower Tarim River desert. The adult plants have a well-developed root system and good drought resistance, salinity tolerance, and adaptability. It is an excellent forage grass, which is important in promoting regional livestock development (Luo et al. 2017 ). However, the morphological establishment stage of A. sparsifolia seedlings is very fragile, particularly under salt and drought stresses. The salt and drought resistance by A. sparsifolia can be achieved via morphological, physiological, and biochemical mechanisms along with other strategies, such as symbiosis with mycorrhizal fungi. Mycorrhizae are important for ecosystem restoration and maintenance of biodiversity (Alessandra et al. 2020 ). Arbuscular mycorrhizal fungi (AMF) colonize plant roots to form mycorrhizal symbionts that complete their life cycle with carbon provided by the plant, whereas the host plant receives the nutrients and water resources that AMF absorbs and transports in a mutually beneficial symbiosis (Debasis et al. 2021). Studies have reported that AMF can enhance the salt and drought tolerance in plants by promoting plant growth, maintaining ion homeostasis, stimulating antioxidant enzyme activity, inducing hormonal signaling responses, and improving photosynthetic capacity (Li et al. 2020 ; Cao et al. 2020 ; Asma et al. 2020 ; Huang et al. 2020 ). AMF transfers nutrients via a close association with the root system, and its growth can extend several centimeters away from the root system to form a dense network of mycelium (Ma et al. 2020 ). Mycelial networks act as a bridge between soil and plant to transport nutrients, such as nitrogen and phosphorus, to host plant roots, thereby promoting plant growth and biomass accumulation (Hu et al. 2020 ; Khirani et al. 2020 ). AMF has been reported to be significantly associated with the desert riparian plant A. sparsifolia , with a colonization frequency of up to 90% and intensity of up to 60% (Yang et al. 2008 ). However, the symbiotic strategy of A. sparsifolia seedlings with AMF under salt and drought stresses is unclear. Therefore, we analyzed the effects of AMF on A. sparsifolia seedlings at the vulnerable growth stage under salt and drought stresses via a multidisciplinary approach. Various physiological and ecological responses of A. sparsifolia seedlings to drought and salt stresses after AMF inoculation were studied in terms of seedling growth, photosynthesis, antioxidant system, osmoregulation, and hormonal response to reveal the AMF-mediated protection mechanisms. The study will help to elucidate the reproductive strategy for the live regeneration of A. sparsifolia in desert riparian forests and will provide insights on the use of AMF as a biological conservation tool in desert ecosystems. Materials And Methods Growth substrate Growth substrate consisted of sand and vermiculite (1:1, V/V). Sand was allowed to pass through a 2-mm sieve to remove impurities and further rinsed under running tap water until the water was clear. Further, the sand was rinsed once with deionized water and sterilized at 121°C for 2 h to eliminate all possible mycorrhizal propagules and other micro-organisms. The sieved and sterilized sand as a growth substrate has the following properties: pH 8.69, organic matter content 1.18 g·kg − 1 , total nitrogen content 0.073 g·kg − 1 , hydrolytic nitrogen content 12 mg·kg − 1 , total phosphorus content 0.505 g·kg − 1 , effective phosphorus content 2.44 mg·kg − 1 , total potassium content 22.9 g·kg − 1 , effective potassium content 82.3 mg·kg − 1 , and conductivity 334 µs·cm − 1 . Vermiculite was autoclaved (121°C, 2 h), mixed well with the prepared sand (1:1, V/V), and used as the growth substrate in this study. Plant And Fungal Inoculum The seeds of A. sparsifolia were collected in August 2020 at the natural habitat restoration demonstration area of the lower Tarim River at the Insu section (40°25.918' N, 87°56.458' E) and were stored at 4°C. Before sowing, the seeds were polished with sandpaper to release dormancy, disinfected with 75% alcohol for 10 min, washed with sterile water, and dried using a filter paper. Claroideoglomus etunicatum and Funneliformis mosseaedominant , the dominant AMF species in the roots of A. sparsifolia , were obtained from Beijing Academy of Agricultural and Forestry Sciences. They were mixed in a ratio of 1:1 (w/w) to prepare the inoculum. The inoculum contained spores (14–20 spores per g inoculum), mycelia, root fragments, and sand. Experimental Design The experiment was conducted in the greenhouse of Xinjiang Normal University, China, with an average temperature of 27°C and a relative air humidity of 30.5%. The experiment consisted of six group of treatments: 1) control/CK-NM (without salt or drought stress and no AMF inoculation), 2) D-NM (drought stress and no AMF inoculation), 3) S-NM (salt stress and no AMF inoculation), 4) CK-AM (AMF inoculation and no salt or drought stress), 5) D-AM (drought stress and AMF inoculation), and 6) S-AM (salt stress and AMF inoculation). Each treatment had six replicates, with a total of 36 plastic pots. Uniformly sized, full-grained A. sparsifolia seeds were selected and sown in plastic pots (24-cm length, 16-cm width, and 18-cm depth). When the seedlings grew to a height of approximately 5 cm, they were transferred to plastic pots sterilized with 75% ethanol. Each plastic pot contained 3 plants and 4 kg of fixed growth substrate. According to the AMF inoculation method, 20 g of inoculum was evenly spread flat at 10 cm from the soil surface. In the control group, equal weight of growth substrate was added instead. During the first 30 days, seedlings were grown without drought or salt stress to obtain plants with functional mycorrhizas and to avoid stress effects on the establishment of symbiosis with AMF. Drought stress was set at 30% ± 5% of the field water holding capacity, and salt stress was set at 0.6% of the weight of growth substrate to keep the A. sparsifolia seedlings under stress but allowing not to die. Soil water content was measured using a WET-2 portable rapid moisture meter. The relative soil water content was adjusted to fall within the range of the drought stress treatment by weighing and rehydrating with an electronic scale at 8:00 pm each day. To avoid osmotic shock, sodium chloride solution was gradually introduced by successively adding 100 mL of prescribed solution of salts in distilled water every 3 days, starting at day 30 after sowing until the sodium chloride content was 0.6% of the weight of the growth substrate. To the groups without salt or drought stress, equal volume of distilled water was added, ensuring that no excess leaching occurred from the pots. A saucer was placed under each pot to retain salt and other nutrients. A total volume of 300 mL of the corresponding salt solution or distilled water was added to each pot. The indicators were measured after 60 days of drought or salt stress. Throughout the experiment, the moisture content of the growth substrate in the groups without drought stress was maintained at 70% ± 5% of the field holding capacity. Mycorrhizal Colonization After harvesting A. sparsifolia , fresh roots were selected and rinsed with tap water. The roots with diameter < 2 mm were further selected and cut into 1-cm-long fragments for fixation, dissociation, acidification, and staining. Before observation and photography, the roots were decolorized with lactic acid and glycerol solution (1:1, V/V), and the stained root samples were squashed with a cover glass onto a glass slide. Mycorrhizal colonization rate and intensity were determined using the Asma method (2020). In total, 20 g soil sample was randomly selected from the root system of A. sparsifolia seedlings, and the number of AMF spores was counted by decantation using a wet sieve. Plant Growth Plant height and basal stem length were measured from the A. sparsifolia seedlings at days 0 and 60 of salt or drought stress. Statistical analyses of root length, root surface area, and root tip number were conducted using WinRHZIO root image analysis software(Zealquest Scientific Technology Co., Ltd, Shanghai, China). The relative growth rates and other parameters were calculated as follows: Growth rate of plant height = (plant height at day 60 − plant height at day 0)/60 × 100% Growth rate of basal stem = (basal stem length at day 60 − basal stem length at day 0)/60 × 100% Specific root length (m/g) = root length/root dry weight Root to shoot ratio = aboveground biomass/belowground biomass Biomass And Nutrient Accumulation During the harvest, the plants were cut from the basal stem, and the above- and belowground parts were divided. Their fresh weight was separately measured. Further, they were placed in a constant-temperature drying oven at 95°C for 15 min to destroy the enzymatic activity in the fresh leaves, followed by drying at 80°C till constant weight was obtained, which was measured using an electronic balance. The nitrogen (N) and phosphorus (P) contents were evaluated at the Institute of Botany, Chinese Academy of Sciences, after drying and grinding the above- and belowground parts. P content (%) was measured using the molybdenum phosphate method, and N content (%) was determined using Kjeldahl method. P content (mg) = %P × weight of biomass N content (mg) = %N × weight of biomass Physiological Measurements The initial fluorescence (Fo), maximum fluorescence (Fm), maximum photochemical efficiency (Fv/Fm), and potential photochemical efficiency (Fv/Fo) of leaves of A. sparsifolia seedlings were measured every 15 days during the treatment period using a MINI-Pam portable modulated chlorophyll fluorometer(Zealquest Scientific Technology Co., Ltd, Shanghai, China). The data were collected from the same seedlings at the same sites on different leaves from the bottom up. The parameters were evaluated using the following formulae. Maximum photochemical efficiency (Fv/Fm) = (Fm − Fo)/Fm Potential photochemical efficiency (Fv/Fo) = (Fm − Fo)/Fo At the harvest, fresh leaf samples (0.1 g) were cleaned using deionized water to remove any surface contamination. Chlorophyll content was determined using the acetone method. The activities of superoxide dismutase (SOD), peroxisome (POD), and catalase (CAT) were determined using the nitrogen blue tetrazolium, guaiacol, and trace methods, respectively. Malondialdehyde (MDA), proline (Pro), and soluble sugar (SS) contents were determined using the thiobarbituric acid, sulfosalicylic acid, and anthrone colorimetric methods, respectively. Each treatment was repeated three times for each physiological index. The kits for determining SOD, POD, and CAT activities and chlorophyll, MDA, Pro, and SS contents were purchased from Beijing Solaibao Technology Co.(Beijing, China) and used as per the manufacturer’s instructions. The levels of hormones [strigolactones (SLs), indole acetic acid (IAA), gibberellic acid (GA), and abscisic acid (ABA)] were measured using enzyme-linked immunosorbent assay (ELISA; EnzymeLink Biologics, Shanghai, China) as per the manufacturer’s instructions. The content of each hormone indicator was evaluated using 3 replicates using a microplate reader. Statistical analysis Data analysis was performed using the Statistical Package for the Social Sciences (SPSS 21.0). Potential differences among various treatments were analyzed using Duncan’s multiple-comparison tests (P < 0.05). All data in the figures and tables are the original data presented as mean ± standard error (mean ± SE). P < 0.05 was considered significant. The results were plotted using origin 2019 and TB tools. Results Mycorrhizal properties The AMF colonization rate and intensity and spore number in the uninoculated groups were zero. The colonization rate of AMF-inoculated seedlings in the CK AM was as high as 87.58%. Salt and drought stresses significantly reduced the AMF colonization rate in the seedlings ( P < 0.05), the rate was slightly higher in the D-AM group than in the S-AM group. AMF colonization intensity was significantly higher in the CK-AM group than in the D-AM group and was significantly higher in the D-AM group than in the S-AM group ( P S-AM > D-AM ( P < 0.05) (Table 1 ). Table 1 Effect of AMF inoculation in the roots of Alhagi sparsifolia seedlings under salt and drought stresses Treatment AMF AMF Colonization rate/(%) AMF Colonization intensity/(%) Number of AMF spores CK NM 0 ± 0c 0 ± 0d 0 ± 0d AM 87.58 ± 2.89a 18.66 ± 0.94c 20.00 ± 0.58a D NM 0 ± 0c 0 ± 0d 0 ± 0d AM 65.34 ± 2.83b 28.97 ± 1.79b 12.00 ± 0.58c S NM 0 ± 0c 0 ± 0d 0 ± 0d AM 62.88 ± 2.44b 48.17 ± 1.62a 13.67 ± 0.33b CK, D, and S represent blank control, drought stress, and salt stress, respectively. NM and AM represent no inoculation and inoculation with AMF, respectively. The results are expressed as the mean ± standard error of three replicates. Duncan’s test revealed that the parameters differed significantly ( P < 0.05) among the treatments. Apparent Seedling Growth AMF inoculation significantly ( P < 0.05) promoted the height and basal stem length in A. sparsifolia seedlings, whereas salt and drought stresses had no significant effect on them. Length, surface area, tip number, and specific length of the root were significantly higher in the AM groups than in the NM groups. The root length of seedlings in the S-AM group was significantly lower than that in the D-AM group ( P < 0.05) (Table 2 ). Under both stress and non-stress conditions, AMF inoculation significantly promoted the growth of above- and belowground parts of A. sparsifolia seedlings; however, the non-stress groups exhibited a stronger promotional effect than the stress groups. Table 2 Effect of AMF inoculation on the height, basal stem diameter, and root growth of A. sparsifolia seedlings under salt and drought stresses Treatment AMF Plant height growth rate/(%) Basal stem growth rate/(%) Root length/(cm) Root surface area/(cm 2 ) Number of root tips Specific root length/(m·g − 1 ) CK NM 2.87 ± 0.41b 0.51 ± 0.12c 13.21 ± 0.11d 3.58 ± 0.05c 6.00 ± 0.58c 1.56 ± 0.03b AM 8.06 ± 0.58a 1.72 ± 0.31a 64.61 ± 0.81a 24.97 ± 1.26a 37.67 ± 3.84a 3.30 ± 0.18a D NM 2.56 ± 0.19b 0.44 ± 0.07c 10.74 ± 0.64d 2.68 ± 0.19c 5.00 ± 0.58c 1.35 ± 0.05b AM 7.61 ± 0.43a 1.36 ± 0.14ab 54.78 ± 0.69b 18.21 ± 1.32b 22.00 ± 2.08b 3.58 ± 0.20a S NM 2.34 ± 0.17b 0.35 ± 0.04c 9.47 ± 0.23d 2.51 ± 0.27c 3.67 ± 0.88c 1.46 ± 0.02b AM 7.44 ± 0.22a 0.88 ± 0.21bc 44.28 ± 0.68c 17.57 ± 1.22b 20.67 ± 3.48b 3.56 ± 0.03a CK, D, and S represent blank control, drought stress, and salt stress, respectively. NM and AM represent no inoculation and inoculation with AMF, respectively. The results are expressed as the mean ± standard error of three replicates. Duncan’s test revealed that the parameters differed significantly ( P < 0.05) between the treatments. Biomass And Root To Shoot Ratio In The Seedlings The fresh and dry weights of above- and belowground parts of A. sparsifolia seedlings increased after AMF inoculation, with the fresh and dry weights of belowground parts increasing significantly ( P < 0.05) (Fig. 1 ). Fresh and dry weights were significantly lower in the AM group than in the CK-AM group under salt and drought stresses, indicating that salt and drought stresses had a negative effect on the inoculated AMF. Among them, the dry weight of the belowground part of the S-AM group was significantly lower than that of the D-AM group, indicating that salt stress had a greater effect on the growth of the belowground part of seedlings than drought stress ( P < 0.05). The root to shoot ratio is an important indicator of plant’s allocation of resources. The root to shoot ratios of dry and fresh weights in the AM group were significantly greater than those of the NM group. This indicated that with the assistance of AMF, A. sparsifolia seedlings allocated more resources for the growth and development of the belowground parts. Compared with the CK-AM group, salt and drought stresses significantly promoted an increase in the root to shoot ratio of fresh weight in the AMF-inoculated groups; however, no significant difference existed between the D-AM and S-AM groups. N And P Contents Salt and drought stresses impair the root growth of A. sparsifolia seedlings, thereby reducing the plant’s uptake and use of N and P elements. The CK-NM group exhibited higher %N and N content than the D-NM and S-NM groups. Both %N and N content were higher in the CK-AM group than in the D-AM and S-AM groups, with significant difference in their N contents ( P < 0.05). The P content exhibited the same trend as the N content; however, the variation was greater, with the AM group exhibiting 5–6-fold higher P content than the NM group. The %P and P content of the D-AM group was significantly higher than that of the S-AM group ( P < 0.05) (Table 3 ), indicating a stronger inhibitory effect of salt stress on access of seedlings to nutrients. Collectively, N and P contents were significantly higher in the no-inoculation group than in the other five groups under salt stress. The N and P contents in the AM group were significantly lower than those in the NM group under salt or drought stress, indicating that AMF inoculation could significantly reduce N and P contents. Table 3 Effect of AMF inoculation on the N and P contents of A. sparsifolia seedlings under salt and drought stresses Treatment AMF N% Nitrogen content/(mg) P% Phosphorus content/(mg) N/P CK NM 1.79 ± 0.01bc 2.56 ± 0.05cd 1.13 ± 0.05d 1.63 ± 0.07d 1.58 ± 0.08c AM 2.22 ± 0.19a 6.01 ± 0.65a 3.50 ± 0.11a 9.48 ± 0.64a 0.63 ± 0.04d D NM 1.71 ± 0.03bc 2.35 ± 0.01cd 0.82 ± 0.01de 1.13 ± 0.01d 2.09 ± 0.02b AM 2.01 ± 0.13ab 3.97 ± 0.30b 3.13 ± 0.07b 6.19 ± 0.37b 0.64 ± 0.06d S NM 1.64 ± 0.01c 1.93 ± 0.04d 0.53 ± 0.01e 0.63 ± 0.00d 3.08 ± 0.04a AM 1.91 ± 0.04abc 3.26 ± 0.16bc 2.54 ± 0.23c 4.35 ± 0.52c 0.76 ± 0.06d CK, D, and S represent blank control, drought stress, and salt stress, respectively. NM and AM represent no inoculation and inoculation with AMF, respectively. The results are expressed as the mean ± standard error of three replicates. Duncan’s test revealed that the parameters differed significantly ( P < 0.05) between the treatments. Chlorophyll Content And Chlorophyll Fluorescence Chlorophyll a, chlorophyll b, chlorophyll content, and chlorophyll a/b exhibited following trend under salt or drought stress and AMF inoculation: CK-AM > CK-NM > D-AM > S-AM > D-NM > S-NM. Therefore, AMF promoted the accumulation of chlorophyll in A. sparsifolia seedlings, and the promotional effect was greater under salt stress than under drought stress. Chlorophyll fluorescence is a rapid, nondestructive method for detecting characteristics of plants under stress and photosynthetic performance. The maximum photochemical efficiency (Fv/Fm) and potential photochemical efficiency (Fv/Fo) of the D-NM and S-NM groups exhibited a significant downward trend with increased duration of salt and drought stresses, with the S-NM group exhibiting the fastest and lowest decrease. The Fv/Fm and Fv/Fo values of the CK-NM and CK-AM groups exhibited a stable trend over time. At day 60, the Fv/Fm and Fv/Fo values of the CK-AM group were significantly higher than those of the D-AM and S-AM groups ( P < 0.05). Comparing the AM and NM groups under salt and drought stresses, respectively, it was clearly observed that AMF inoculation significantly alleviated the effects of salt and drought stresses on the Fv/Fm and Fv/Fo values of the seedlings, particularly in the S-AM group (Fig. 2). Figure 2 Effect of AMF inoculation on chlorophyll content and chlorophyll fluorescence in A. sparsifolia seedlings under salt and drought stresses. CK, D, and S represent blank control, drought stress, and salt stress, respectively. NM and AM represent no inoculation and inoculation with AMF, respectively. The results are expressed as the mean ± standard error of three replicates. Duncan’s test revealed that the parameters differed significantly ( P < 0.05) between the treatments. Activities of antioxidant enzymes Figure 3 shows that both salt or drought stress and AMF inoculation significantly increased the activities of antioxidant enzymes SOD, POD, and CAT in A. sparsifolia seedlings ( P < 0.05). The S-AM group exhibited the highest antioxidant enzyme activity. SOD activity was significantly higher in the AM group than in the NM group under salt and drought stresses. It was significantly higher in the S-AM group than in the D-AM group, with no significant difference between AM and NM in the CK group. POD activity was significantly higher in the AM group than in the NM group; it was significantly higher under salt stress than under drought stress and was significantly higher under drought stress than in the blank control. The trend of CAT activity was the same as that of POD. This indicated that AMF inoculation increased the antioxidant enzyme activity in the seedlings in response to the damage caused by salt and drought stresses. Figure 3 Effect of AMF inoculation on the activities of antioxidant enzymes (SOD, POD, and CAT) in A. sparsifolia seedlings under salt and drought stresses. CK, D, and S represent blank control, drought stress, and salt stress, respectively. NM and AM represent no inoculation and inoculation with AMF, respectively. The results are expressed as the mean ± standard error of three replicates. Duncan’s test revealed that the parameters differed significantly ( P < 0.05) between the treatments. Mda, Pro, And Ss Contents MDA, Pro, and SS contents increased in A. sparsifolia seedlings under salt and drought stresses (Fig. 4 ). The MDA content decreased significantly after AMF inoculation, exhibiting a trend CK-AM < D-AM < S-AM. Pro and SS contents significantly increased after AMF inoculation, and they were significantly higher under salt stress than under drought stress ( P < 0.05). Therefore, AMF could protect itself from salt and drought stresses to some extent by assisting A. sparsifolia seedlings to accumulate osmoregulatory substances. Hormonal Levels Salt and drought stresses significantly increased the IAA and ABA levels, reduced the GA level ( P 0.05) in A. sparsifolia seedlings (Fig. 5 ). After AMF inoculation, IAA level significantly increased with its level significantly lower in the S-AM group than in the D-AM group; ABA level significantly decreased with its level significantly higher in the S-AM group than in the D-AM group. The GA level was significantly higher in the S-AM group than in the S-NM group but was significantly lower in the D-AM group than in the D-NM group. This indicated that drought and salt stresses regulated the GA level of AMF-inoculated seedlings via various ways. The SLs level was significantly reduced after AMF inoculation, with no significant difference between the salt- and drought-stress groups. Correlation Analysis The heat map and Table 4 show the correlation between various indicators of growth and physiology of A. sparsifolia seedlings under the six treatments, namely, CK-NM, D-NM, S-NM, CK-AM, D-AM, and S-AM (Fig. 6 ; Table 4 ). The CK-NM, D-NM, and S-NM groups exhibited negative correlations for morphological growth and most physiological indicators and positive correlations for ABA and MDA levels, particularly under salt stress. The positive correlation among growth, biomass accumulation, and chlorophyll content in the above- and belowground parts of seedlings in the CK-AM group was very strong after AMF inoculation. This indicated that AMF inoculation significantly promoted the growth of A. sparsifolia seedlings under non-stress conditions. AMF inoculation alleviated salt- and drought-stress-induced damage to seedlings, which was highlighted by the increased antioxidant enzyme activity and activated osmoregulatory systems of seedlings under salt stress. Antioxidant enzyme activity and osmoregulatory substance content exhibited a significant positive correlation in the S-AM group. Table 4 Effect of salt or drought stress, AMF inoculation, and their interactions on all measured response variables tested using multivariate ANOVA indicators Drought Salt AMF Drought*AMF Salt*AMF AMF Colonization rate 33.279*** 41.037*** 1769.429*** 33.279*** 41.037*** AMF Colonization intensity 6.032* 49.421*** 377.183*** 6.032* 49.421*** Number of AMF spores 110.979*** 66.083*** 1734.289*** 110.979*** 66.083*** Plant height growth rate 0.194ns 0.446ns 49.299*** 0.007ns 0.002ns Basal stem growth rate 1.286ns 6.984* 26.973*** 0.719ns 3.449ns Root length 12.755** 17.163** 101.362*** 9.878** 9.878** Root surface area 5.520* 21.143** 354.611*** 1.976ns 10.043** Number of root tips 17.908** 21.942** 461.500*** 10.489** 12.172** Specific root length 0.146ns 0.518ns 464.981*** 4.723* 2.469ns Fresh weight above ground 19.456** 27.574*** 15.686** 5.338* 3.574ns Fresh weight below ground 9.000* 10.562** 108.372*** 0ns 0.174ns Dry weight above ground 8.000* 18.00** 7.111* 0ns 0.500ns Dry weight below ground 19.853** 74.206*** 250.373*** 10.676** 19.853** Root to shoot ratio(DW) 0.536ns 1.051ns 56.146*** 1.737ns 1.678ns Root to shoot ratio(FW) 0.106ns 0.854ns 177.787*** 17.006** 10.349** %N 2.192ns 5.481* 14.546** 0.523ns 0.683ns N content 13.777** 31.152*** 52.944*** 9.114* 12.193** %P 9.783** 50.434*** 552.846*** 0.082ns 2.663ns P content 26.182*** 68.614*** 261.854*** 14.247** 31.145*** N/P 24.751*** 246.347*** 1477.015*** 22.872*** 174.026*** Chlorophyll a concentration 218.337*** 672.622*** 261.534*** 28.105*** 38.587*** Chlorophyll b concentration 68.948*** 110.639*** 40.615*** 19.144** 5.868* Chlorophyll contentration 18.678** 164.56*** 81.694*** 77.101*** 77.101*** Chlorophyll a/b ratio 249.785*** 649.343*** 248.531*** 2.412ns 11.139** Fv/Fm 13.135** 49.662*** 0.273ns 0.041ns 4.054ns Fv/Fo 13.374** 37.668*** 0.900ns 0.308ns 3.115ns Superoxide dismutase SOD 23.630*** 110.041*** 92.768*** 6.692* 22.075** Peroxidase POD 16.071** 79.117*** 26.504*** 0.294ns 0.066ns Catalase CAT 29.390*** 220.195*** 240.587*** 0.691ns 45.789*** Malondialdehyde MDA 73.554*** 295.275*** 111.881*** 0.250ns 0.324ns Proline Pro 103.597*** 1255.071*** 1287.975*** 15.402** 219.118*** Soluble sugar SS 5.960* 26.601*** 63.209*** 1.136ns 5.467* Indoleacetic acid IAA 0.434ns 7.261* 2715.657*** 101.609*** 456.749*** Gibberellin GA 67.372*** 287.883*** 23.061*** 28.138*** 154.547*** Abscisic acid ABA 611.830*** 1304.326*** 220.300*** 60.476*** 43.074*** Strigolactones SLs 2.247ns 9.867** 50.724*** 0.044ns 0.147ns F-values are followed by P -values ns P ༞0.05 * P < 0.05 ** P < 0.01 *** P < 0.001 Discussion A. sparsifolia is an ecologically important plant. The adult plants are ideal for ecological restoration in arid areas. However, salt and drought stresses endanger the morphological establishment of seedlings, which is a key stage in the growth of A. sparsifolia . Therefore, these stresses limit the process of live regeneration of A. sparsifolia , and the stability of the population could not be maintained. In this study, we employed the natural environment of the Tarim River basin with a soil salt content of 0.41–6.28% (Zhong et al. 2018 ) and the salt tolerance range of 0.4–0.8% for A. sparsifolia seed germination (Zhang et al. 2009). Based on this condition and our previous study (Zhang et al. 2016; Zainur 2021; GAO et al. 2022 ), we conducted this study under salt stress (0.6%) and drought stress (30% ± 5%) to verify whether AMF can improve the salt and drought tolerance of A. sparsifolia seedlings and to reveal the survival strategies during the seedling period by altering apparent growth, photosynthetic efficiency, antioxidant enzyme activity, and levels of osmoregulatory substances and hormones. Effects Of Amf Inoculation On Salt- Or Drought-induced Damage In The Seedlings Studies have reported that the root system of A. sparsifolia seedlings can establish a good symbiosis with AMF, which is a highly clumped mycorrhizal-dependent plant (Wang et al. 2010 ). The rate and intensity of AMF colonization were significantly reduced under salt and drought stresses, indicating that salt and drought stresses significantly inhibited the growth of AMF. The present study reported that although the colonization intensity of AMF under salt stress was significantly lower than that under drought stress, the spore number of AMF was significantly higher under salt stress than under drought stress. This suggested that the massive accumulation of AMF spores may be an effective reproductive strategy for its successful survival in adverse conditions. Salt and drought stresses inhibited the growth and live regeneration processes in the seedlings (Table 4 ); however, AMF promoted host plant growth by expanding the total root uptake area and improving uptake of nutrients by the plant (Diagne et al. 2020 ). The height and basal stem length of A. sparsifolia seedlings significantly increased after AMF inoculation under both stress and non-stress conditions. After AMF inoculation, the root length increased by 3.8, 4.1, and 3.6-fold in the non-stressed, drought-stressed, and salt-stressed groups, respectively. The specific root length and root to shoot ratio increased by 1 fold in the D-AM groups, indicating that AMF responded to the damage due to drought stress by enhancing the root growth of the seedlings. Adaptation in the form of changes in root morphology and microbial cooperation require the allocation of carbon from photosynthesis to compete with the belowground carbon sinks, either to promote root growth or to transfer to symbiotic partners (Cao et al. 2020 ). Plant biomass allocation is the result of a combination of biotic and abiotic factors, and the exponential increase in the root to shoot ratio of the seedlings under abiotic and non-stress factors confirmed the significant promotion of root growth by the biotic factor AMF. The exponential increase in the specific root length further confirmed the beneficial effect of AMF on the increase of root length during the root growth of the seedlings. Effects Of Amf Inoculation On N And P Contents Under Salt Or Drought Stress N and P are the two elements that are essential for plant growth. Notably, AMF inoculation significantly promoted the uptake of elemental P. The low fixation of P in soil and its rapid precipitation with soil minerals result in low effective P content in soil, whereas AMF contributes to 80% of total P uptake and has the potential to secrete acid phosphatase and organic acids that directly promote P uptake (Masao et al. 2020 ; Song et al. 2021 ). The importance of AMF in P uptake was demonstrated by the 4–5-fold increase in the P uptake after AMF inoculation in this study. The contribution of AMF to N uptake is usually smaller than that to P uptake by plants; however, AMF can still transfer N to the host (Wang et al. 2022 ). In this study, the N content in the AMF-inoculated groups was higher than that in the non-inoculated groups and was more influenced by stress factors. The AMF-inoculated seedlings had the highest total N content under non-stress conditions. The AMF-inoculated seedlings had a higher total N content than the non-inoculated seedlings under salt and drought stresses, and the non -inoculated seedlings had the lowest total N content under salt stress. Both salt and drought stresses and AMF inoculation exhibited important effects on the growth of seedlings. N:P is an accurate indicator of the nutrient limitation for plant growth. N:P of A. sparsifolia seedlings obtained in this study ranged from 0.6 to 3.1; this indicated that AMF helped the seedlings in increasing their P content while leaving them relatively deficient in N, a phenomenon that was more pronounced under salt and drought stresses. Effects Of Amf Inoculation On The Photosynthesis Under Salt Or Drought Stress Chlorophyll content is an indicator of photosynthetic status, developmental stage, and degree of stress (Li et al. 2020 ). The chlorophyll content in A. sparsifolia seedlings under salt stress was significantly lower than that under drought stress and was significantly lower in the stress groups than in the non-stress groups. This indicated that both salt and drought stresses can impair the growth of seedlings by reducing their chlorophyll content, and in turn photosynthetic capacity; however, this effect was greater under salt stress. After AMF inoculation, the chlorophyll content of seedlings significantly increased, indicating that the presence of AMF and salt stress had less effect on chlorophyll synthesis. This could be attributed to the substances secreted by AMF, such as cytokinins, as these substances facilitate chloroplast development and increase chlorophyll levels (Wang et al. 2022 ). Photosystem II (PSII) is the main light-driven reaction involved in photosynthesis (Wu et al. 2020 ). Salt and drought stresses reduced the photochemical efficiency and electron transfer in the PSII reaction center, resulting in lower Fv/Fm and Fv/Fo values than normal in the leaves of A. sparsifolia seedlings. However, Fv/Fm and Fv/Fo values did not significantly decrease with increasing duration of stress after AMF inoculation. This indicated that salt and drought stresses had a negative impact on the photosynthetic physiology of the seedlings, and salt stress had a greater impact; AMF inoculation had a sustained but not significant effect in alleviating the effects of the stresses. Effects Of Amf Inoculation On The Antioxidant Enzyme Activities Under Salt Or Drought Stress AMF promotes plant growth, increases the above- and belowground biomass, and mitigates the negative effects of abiotic stresses (Asma et al. 2020 ; Dorotéia et al. 2020 ; Wang et al. 2020 ). AMF inoculation activated the antioxidant enzyme system of A. sparsifolia seedlings and increased the activity of SOD, POD, and CAT, which was particularly significant under salt stress compared with that under drought stress. The antioxidant enzyme activity was most correlated with AMF inoculation in terms of physiological indicators in the seedlings. This could be because of following reasons. First, salt stress increased the accumulation of toxic reactive oxygen species (ROS) and stimulated the seedlings’ own antioxidant defense system to produce numerous antioxidant enzymes. Second, AMF protected the seedlings from ROS by enhancing antioxidant enzyme activity to maintain an effective antioxidant enzyme system. Furthermore, numerous studies have reported that the antioxidant enzyme activity of AMF-inoculated plants is higher than that of non-inoculated plants in this species (Wang et al. 2020 ). The antioxidant enzyme activity was increased under drought stress as well; however, the effect was lower than that under salt stress, presumably because the levels of osmotic pressure and osmoregulatory substances in the seedlings were greater under salt stress than under drought stress. Effects Of Amf Inoculation On Mda, Pro, And Ss Contents Under Salt Or Drought Stress Salt stress can directly lead to osmotic stress, primary ion toxicity, and secondary stresses such as nutrient deficiency and oxidative stress in plants, which ultimately hinder plant growth (Yang et al. 2019 ). The effects of drought stress are similar to those of salt stress. Both salt and drought stresses disrupt cell structure and some key physiological functions in plants, e.g., cell membrane damage, reduced photosynthetic capacity, production of ROS, and accumulation of harmful substances (Li et al. 2020 ). The results of this study demonstrated that the content of osmoregulatory substances increased in A. sparsifolia seedlings, and the increase was higher under salt stress. Pro is the main store of energy and N under salt stress (Wang et al. 2020 ). After AMF inoculation, the Pro content increased significantly under salt stress. Therefore, AMF helped the plant in accumulating more Pro and overcoming the salt-stress-induced osmotic imbalance. Effects Of Amf Inoculation On Hormonal Levels Under Salt Or Drought Stress AMF not only improves macroscopic plant growth and N and P uptake but also regulates microscopic intracellular hormonal levels. AMF colonization relies on the release of SLs into the root zone to germinate spores and attract mycelium to the root system (Ma et al. 2020 ). In this study, the level of SLs in the AMF-inoculated groups was significantly lower than that in the non-inoculated groups. It was assumed that SLs were released into the soil for the colonization of AMF with the root system in the AMF-inoculated group. SLs can regulate the process of adaptation of plants to adverse conditions (Liao et al. 2016 ). In this study, the level of SLs was significantly lower in the non-inoculated group under salt stress than in the blank control group. Compared with the blank control group, the increase in the IAA and ABA levels and decrease in the GA level under salt and drought stresses demonstrated the self-regulation ability of the seedlings under salt and drought stresses. The significant increase in the IAA level and significant decrease in the GA level in this study supported the idea that significant changes in hormonal levels are also a survival strategy for seedlings to mitigate the effects of salt and drought stresses with the help of AMF (Huang et al. 2020 ). AMF enhanced the tolerance of the seedlings to salt and drought stresses through a combination of mechanisms, reaffirming the critical role of mycorrhizal fungi as a major component of terrestrial ecosystems, particularly in the restoration of degraded desert ecosystems (Meng et al. 2008 ). Conclusion In summary, we can conclude that A. sparsifolia –AMF symbiosis mitigated the negative effects of salt and drought stresses on A. sparsifolia seedlings by altering plant growth and physiology. The mechanisms are as follows. AMF inoculation promoted the growth of above- and belowground parts, facilitating more nutrient gain and biomass accumulation. It better balanced light energy distribution in photochemical and non-photochemical processes. It increased antioxidant enzyme activity to scavenge and reduce the levels of ROS. It increased the levels of soluble substances to improve osmoregulation. It up- or downregulated hormonal levels to induce hormone signaling responses, ultimately resulting in better salt and drought tolerance by the seedlings. Therefore, AMF has important potential application in the regeneration of A. sparsifolia and restoration of degraded desert ecosystems in arid zones. Declarations Acknowledgments This work was financially supported by the National Natural Science Foundation of China (No. 42067067) and the Natural Science Foundation of Xinjiang Uygur Autonomous Region (No.2020D01A74). Thanks to Dr. Jingbo Zhang for guidance in TB tools data processing software. Author Contributions Formal analysis: Xiaodong Ma,Aili Yilinuer. Resources:Dawuti Maigepiretiguli. Software: Haiou Wang. Supervision: Xiaonan Chen. Writing - original draft: Xiaonan Chen. Writing - review: Xiaodong Ma. Declarations Conflict of interest The author declared that they have no conflict of interest. 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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-2374175","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":160686148,"identity":"294cfbb9-84e7-447d-9f94-9a794fbd958c","order_by":0,"name":"Xiaonan Chen","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA2ElEQVRIiWNgGAWjYBACfvnzHx///SPBw8/MfIA4LZIzGIwNeBssZCTb2xKI02Jwg8FMgrehwsbgzBkDIl12uyHZQHKHBI/BjZyPN94w2MnpNhDQwTjnwMEHhmckeCRv5G62nMOQbGx2gIAWZobEZoMENgkevhu526R5GA4kbiOkhY0hmU3iAFALw42cZ8Rp4ZFIY5NsbJPgEThzho04LRI8Z5iNGUB+aW8ztpxjQIRf7I/3MD5mqKizB0blwxtvKuzkCGpBs5LYqEHSQqqOUTAKRsEoGBEAALRAQBwVYEeYAAAAAElFTkSuQmCC","orcid":"","institution":"Xinjiang Normal University","correspondingAuthor":true,"submittingAuthor":false,"prefix":"","firstName":"Xiaonan","middleName":"","lastName":"Chen","suffix":""},{"id":160686149,"identity":"4f2972f3-20f6-49cd-b85f-09ce1b568dfe","order_by":1,"name":"Aili Yilinuer","email":"","orcid":"","institution":"Xinjiang Normal University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Aili","middleName":"","lastName":"Yilinuer","suffix":""},{"id":160686150,"identity":"c2be76d9-a1d2-40ff-ae19-e9b65e076744","order_by":2,"name":"Xiaodong Ma","email":"","orcid":"","institution":"Xinjiang Normal University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Xiaodong","middleName":"","lastName":"Ma","suffix":""},{"id":160686151,"identity":"48f4f74b-9ad7-4439-baea-e2888fd5d5c2","order_by":3,"name":"Haiou Wang","email":"","orcid":"","institution":"Xinjiang Normal University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Haiou","middleName":"","lastName":"Wang","suffix":""},{"id":160686152,"identity":"768457a6-d816-4d1e-a286-7864a9f83c1b","order_by":4,"name":"Dawuti Maigepiretiguli","email":"","orcid":"","institution":"Xinjiang Normal University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Dawuti","middleName":"","lastName":"Maigepiretiguli","suffix":""}],"badges":[],"createdAt":"2022-12-13 13:31:58","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-2374175/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-2374175/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s10725-023-00996-0","type":"published","date":"2023-04-05T20:22:47+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":30586808,"identity":"8c6483fa-42dd-43e6-825c-47fc469a55e4","added_by":"auto","created_at":"2022-12-20 19:22:32","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":109193,"visible":true,"origin":"","legend":"\u003cp\u003eEffect of AMF inoculation on the fresh and dry weights of above- and belowground parts and root to shoot ratio of \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings under salt and drought stresses. CK, D, and S represent blank control, drought stress, and salt stress, respectively. NM and AM represent no inoculation and inoculation with AMF, respectively. The results are expressed as the mean ± standard error of three replicates. Duncan’s test revealed that the parameters differed significantly (\u003cem\u003eP\u003c/em\u003e\u0026lt; 0.05) between the treatments.\u003c/p\u003e","description":"","filename":"Fig.1.png","url":"https://assets-eu.researchsquare.com/files/rs-2374175/v1/e153b523ed714a8493bd338c.png"},{"id":30586811,"identity":"52777112-8a39-4852-9c5b-5689f3b8d954","added_by":"auto","created_at":"2022-12-20 19:22:32","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":142022,"visible":true,"origin":"","legend":"\u003cp\u003eEffect of AMF inoculation on chlorophyll content and chlorophyll fluorescence in \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings under salt and drought stresses. CK, D, and S represent blank control, drought stress, and salt stress, respectively. NM and AM represent no inoculation and inoculation with AMF, respectively. The results are expressed as the mean ± standard error of three replicates. Duncan’s test revealed that the parameters differed significantly (\u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05) between the treatments.\u003c/p\u003e","description":"","filename":"Fig.2.png","url":"https://assets-eu.researchsquare.com/files/rs-2374175/v1/2e723928a2865668ca39aff8.png"},{"id":30587804,"identity":"173c30d1-5aad-46af-bddc-e947ac3f74a9","added_by":"auto","created_at":"2022-12-20 19:30:32","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":54467,"visible":true,"origin":"","legend":"\u003cp\u003eEffect of AMF inoculation on the activities of antioxidant enzymes (SOD, POD, and CAT) in \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings under salt and drought stresses. CK, D, and S represent blank control, drought stress, and salt stress, respectively. NM and AM represent no inoculation and inoculation with AMF, respectively. The results are expressed as the mean ± standard error of three replicates. Duncan’s test revealed that the parameters differed significantly (\u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05) between the treatments.\u003c/p\u003e","description":"","filename":"Fig.3.png","url":"https://assets-eu.researchsquare.com/files/rs-2374175/v1/e6de062cd5dbd6c879694af8.png"},{"id":30588125,"identity":"fce434b5-930a-4638-be50-2926448cbd09","added_by":"auto","created_at":"2022-12-20 19:38:32","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":53721,"visible":true,"origin":"","legend":"\u003cp\u003eEffect of AMF inoculation on osmoregulatory substances (MDA, Pro, and SS) in \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings under salt and drought stresses. CK, D, and S represent blank control, drought stress, and salt stress, respectively. NM and AM represent no inoculation and inoculation with AMF, respectively. The results are expressed as the mean ± standard error of three replicates. Duncan’s test revealed that the parameters differed significantly (\u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05) between the treatments.\u003c/p\u003e","description":"","filename":"Fig.4.png","url":"https://assets-eu.researchsquare.com/files/rs-2374175/v1/a4d54b96afababacedeea398.png"},{"id":30587805,"identity":"64a7d399-1fd3-46f3-8798-c47a484000c8","added_by":"auto","created_at":"2022-12-20 19:30:32","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":76601,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003e5\u003c/strong\u003e Effect of AMF inoculation on the level of hormones (IAA, GA, ABA, and SLs) in \u003cem\u003eA. sparsifolia \u003c/em\u003eseedlings under salt and drought stresses.\u003cbr\u003e\n CK, D, and S represent blank control, drought stress, and salt stress, respectively. NM and AM represent no inoculation and inoculation with AMF, respectively. The results are expressed as the mean ± standard error of three replicates. Duncan’s test revealed that the parameters differed significantly (\u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05) between the treatments.\u003c/p\u003e","description":"","filename":"Fig.5.png","url":"https://assets-eu.researchsquare.com/files/rs-2374175/v1/0c0968af90f081126ebc3171.png"},{"id":30586813,"identity":"573f489b-d1b3-4ab3-9223-4c35238780a8","added_by":"auto","created_at":"2022-12-20 19:22:32","extension":"jpg","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":859241,"visible":true,"origin":"","legend":"\u003cp\u003eHeatmap of the combined effects of AMF inoculation on various growth and physiological indicators of \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings under salt and drought stresses. CK-NM, D-NM, S-NM, CK-AM, D-AM, and S-AM represent the no salt and drought stresses and no AMF inoculation control, drought stress and no AMF inoculation, salt stress and no AMF inoculation, no salt and drought stresses and AMF inoculation, drought stress and AMF inoculation, and salt stress and AMF inoculation, respectively.\u003c/p\u003e","description":"","filename":"Fig.6.jpg","url":"https://assets-eu.researchsquare.com/files/rs-2374175/v1/3ed49e3ac6d962f987c899c4.jpg"},{"id":44724254,"identity":"2ae5b6f6-8e4f-4cee-8554-35397735e6e1","added_by":"auto","created_at":"2023-10-16 20:28:05","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1079633,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-2374175/v1/e94c416b-3425-4464-8db0-d19954a4bd79.pdf"}],"financialInterests":"","formattedTitle":"Mycorrhizal fungal colonization promotes apparent growth and physiology of Alhagi sparsifolia seedlings under salt or drought stress at vulnerable developmental stage","fulltext":[{"header":"Introduction","content":"\u003cp\u003eSalt and drought stresses are some of the most frequent and severe abiotic stresses in desert ecosystems. They negatively affect many physiological and biochemical indicators and growth and development in plants (Abderrahim et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The lower Tarim River is a typical damaged desert ecosystem in the arid zone, and artificial water transfer projects have been implemented for 22 years to solve the problem of ecological water shortage. Downstream desert riparian forests are located in extremely arid inland areas, often with high salt accumulation in their artificially delivered surface and groundwater. This causes severe stress to the plant seedlings for their survival and growth (Zhang et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Chen et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Li et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). \u003cem\u003eA. sparsifolia\u003c/em\u003e is a perennial herb in the legume family and a dominant herb in the riparian forests of the lower Tarim River desert. The adult plants have a well-developed root system and good drought resistance, salinity tolerance, and adaptability. It is an excellent forage grass, which is important in promoting regional livestock development (Luo et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). However, the morphological establishment stage of \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings is very fragile, particularly under salt and drought stresses. The salt and drought resistance by \u003cem\u003eA. sparsifolia\u003c/em\u003e can be achieved via morphological, physiological, and biochemical mechanisms along with other strategies, such as symbiosis with mycorrhizal fungi.\u003c/p\u003e \u003cp\u003eMycorrhizae are important for ecosystem restoration and maintenance of biodiversity (Alessandra et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Arbuscular mycorrhizal fungi (AMF) colonize plant roots to form mycorrhizal symbionts that complete their life cycle with carbon provided by the plant, whereas the host plant receives the nutrients and water resources that AMF absorbs and transports in a mutually beneficial symbiosis (Debasis et al. 2021). Studies have reported that AMF can enhance the salt and drought tolerance in plants by promoting plant growth, maintaining ion homeostasis, stimulating antioxidant enzyme activity, inducing hormonal signaling responses, and improving photosynthetic capacity (Li et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Cao et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Asma et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Huang et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). AMF transfers nutrients via a close association with the root system, and its growth can extend several centimeters away from the root system to form a dense network of mycelium (Ma et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Mycelial networks act as a bridge between soil and plant to transport nutrients, such as nitrogen and phosphorus, to host plant roots, thereby promoting plant growth and biomass accumulation (Hu et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Khirani et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). AMF has been reported to be significantly associated with the desert riparian plant \u003cem\u003eA. sparsifolia\u003c/em\u003e, with a colonization frequency of up to 90% and intensity of up to 60% (Yang et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). However, the symbiotic strategy of \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings with AMF under salt and drought stresses is unclear.\u003c/p\u003e \u003cp\u003eTherefore, we analyzed the effects of AMF on \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings at the vulnerable growth stage under salt and drought stresses via a multidisciplinary approach. Various physiological and ecological responses of \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings to drought and salt stresses after AMF inoculation were studied in terms of seedling growth, photosynthesis, antioxidant system, osmoregulation, and hormonal response to reveal the AMF-mediated protection mechanisms. The study will help to elucidate the reproductive strategy for the live regeneration of \u003cem\u003eA. sparsifolia\u003c/em\u003e in desert riparian forests and will provide insights on the use of AMF as a biological conservation tool in desert ecosystems.\u003c/p\u003e"},{"header":"Materials And Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eGrowth substrate\u003c/h2\u003e \u003cp\u003eGrowth substrate consisted of sand and vermiculite (1:1, V/V). Sand was allowed to pass through a 2-mm sieve to remove impurities and further rinsed under running tap water until the water was clear. Further, the sand was rinsed once with deionized water and sterilized at 121\u0026deg;C for 2 h to eliminate all possible mycorrhizal propagules and other micro-organisms. The sieved and sterilized sand as a growth substrate has the following properties: pH 8.69, organic matter content 1.18 g\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, total nitrogen content 0.073 g\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, hydrolytic nitrogen content 12 mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, total phosphorus content 0.505 g\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, effective phosphorus content 2.44 mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, total potassium content 22.9 g\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, effective potassium content 82.3 mg\u0026middot;kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, and conductivity 334 \u0026micro;s\u0026middot;cm\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e. Vermiculite was autoclaved (121\u0026deg;C, 2 h), mixed well with the prepared sand (1:1, V/V), and used as the growth substrate in this study.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003ePlant And Fungal Inoculum\u003c/h3\u003e\n\u003cp\u003eThe seeds of \u003cem\u003eA. sparsifolia\u003c/em\u003e were collected in August 2020 at the natural habitat restoration demonstration area of the lower Tarim River at the Insu section (40\u0026deg;25.918' N, 87\u0026deg;56.458' E) and were stored at 4\u0026deg;C. Before sowing, the seeds were polished with sandpaper to release dormancy, disinfected with 75% alcohol for 10 min, washed with sterile water, and dried using a filter paper.\u003c/p\u003e \u003cp\u003e \u003cem\u003eClaroideoglomus etunicatum\u003c/em\u003e and \u003cem\u003eFunneliformis mosseaedominant\u003c/em\u003e, the dominant AMF species in the roots of \u003cem\u003eA. sparsifolia\u003c/em\u003e, were obtained from Beijing Academy of Agricultural and Forestry Sciences. They were mixed in a ratio of 1:1 (w/w) to prepare the inoculum. The inoculum contained spores (14\u0026ndash;20 spores per g inoculum), mycelia, root fragments, and sand.\u003c/p\u003e\n\u003ch3\u003eExperimental Design\u003c/h3\u003e\n\u003cp\u003eThe experiment was conducted in the greenhouse of Xinjiang Normal University, China, with an average temperature of 27\u0026deg;C and a relative air humidity of 30.5%. The experiment consisted of six group of treatments: 1) control/CK-NM (without salt or drought stress and no AMF inoculation), 2) D-NM (drought stress and no AMF inoculation), 3) S-NM (salt stress and no AMF inoculation), 4) CK-AM (AMF inoculation and no salt or drought stress), 5) D-AM (drought stress and AMF inoculation), and 6) S-AM (salt stress and AMF inoculation). Each treatment had six replicates, with a total of 36 plastic pots.\u003c/p\u003e \u003cp\u003eUniformly sized, full-grained \u003cem\u003eA. sparsifolia\u003c/em\u003e seeds were selected and sown in plastic pots (24-cm length, 16-cm width, and 18-cm depth). When the seedlings grew to a height of approximately 5 cm, they were transferred to plastic pots sterilized with 75% ethanol. Each plastic pot contained 3 plants and 4 kg of fixed growth substrate. According to the AMF inoculation method, 20 g of inoculum was evenly spread flat at 10 cm from the soil surface. In the control group, equal weight of growth substrate was added instead. During the first 30 days, seedlings were grown without drought or salt stress to obtain plants with functional mycorrhizas and to avoid stress effects on the establishment of symbiosis with AMF. Drought stress was set at 30% \u0026plusmn; 5% of the field water holding capacity, and salt stress was set at 0.6% of the weight of growth substrate to keep the \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings under stress but allowing not to die. Soil water content was measured using a WET-2 portable rapid moisture meter. The relative soil water content was adjusted to fall within the range of the drought stress treatment by weighing and rehydrating with an electronic scale at 8:00 pm each day. To avoid osmotic shock, sodium chloride solution was gradually introduced by successively adding 100 mL of prescribed solution of salts in distilled water every 3 days, starting at day 30 after sowing until the sodium chloride content was 0.6% of the weight of the growth substrate. To the groups without salt or drought stress, equal volume of distilled water was added, ensuring that no excess leaching occurred from the pots. A saucer was placed under each pot to retain salt and other nutrients. A total volume of 300 mL of the corresponding salt solution or distilled water was added to each pot. The indicators were measured after 60 days of drought or salt stress. Throughout the experiment, the moisture content of the growth substrate in the groups without drought stress was maintained at 70% \u0026plusmn; 5% of the field holding capacity.\u003c/p\u003e\n\u003ch3\u003eMycorrhizal Colonization\u003c/h3\u003e\n\u003cp\u003eAfter harvesting \u003cem\u003eA. sparsifolia\u003c/em\u003e, fresh roots were selected and rinsed with tap water. The roots with diameter\u0026thinsp;\u0026lt;\u0026thinsp;2 mm were further selected and cut into 1-cm-long fragments for fixation, dissociation, acidification, and staining. Before observation and photography, the roots were decolorized with lactic acid and glycerol solution (1:1, V/V), and the stained root samples were squashed with a cover glass onto a glass slide. Mycorrhizal colonization rate and intensity were determined using the Asma method (2020). In total, 20 g soil sample was randomly selected from the root system of \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings, and the number of AMF spores was counted by decantation using a wet sieve.\u003c/p\u003e\n\u003ch3\u003ePlant Growth\u003c/h3\u003e\n\u003cp\u003ePlant height and basal stem length were measured from the \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings at days 0 and 60 of salt or drought stress. Statistical analyses of root length, root surface area, and root tip number were conducted using WinRHZIO root image analysis software(Zealquest Scientific Technology Co., Ltd, Shanghai, China). The relative growth rates and other parameters were calculated as follows:\u003c/p\u003e \u003cp\u003eGrowth rate of plant height = (plant height at day 60\u0026thinsp;\u0026minus;\u0026thinsp;plant height at day 0)/60 \u0026times; 100%\u003c/p\u003e \u003cp\u003eGrowth rate of basal stem = (basal stem length at day 60\u0026thinsp;\u0026minus;\u0026thinsp;basal stem length at day 0)/60 \u0026times; 100%\u003c/p\u003e \u003cp\u003eSpecific root length (m/g)\u0026thinsp;=\u0026thinsp;root length/root dry weight\u003c/p\u003e \u003cp\u003eRoot to shoot ratio\u0026thinsp;=\u0026thinsp;aboveground biomass/belowground biomass\u003c/p\u003e\n\u003ch3\u003eBiomass And Nutrient Accumulation\u003c/h3\u003e\n\u003cp\u003eDuring the harvest, the plants were cut from the basal stem, and the above- and belowground parts were divided. Their fresh weight was separately measured. Further, they were placed in a constant-temperature drying oven at 95\u0026deg;C for 15 min to destroy the enzymatic activity in the fresh leaves, followed by drying at 80\u0026deg;C till constant weight was obtained, which was measured using an electronic balance. The nitrogen (N) and phosphorus (P) contents were evaluated at the Institute of Botany, Chinese Academy of Sciences, after drying and grinding the above- and belowground parts. P content (%) was measured using the molybdenum phosphate method, and N content (%) was determined using Kjeldahl method.\u003c/p\u003e \u003cp\u003eP content (mg) = %P \u0026times; weight of biomass\u003c/p\u003e \u003cp\u003eN content (mg) = %N \u0026times; weight of biomass\u003c/p\u003e\n\u003ch3\u003ePhysiological Measurements\u003c/h3\u003e\n\u003cp\u003eThe initial fluorescence (Fo), maximum fluorescence (Fm), maximum photochemical efficiency (Fv/Fm), and potential photochemical efficiency (Fv/Fo) of leaves of \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings were measured every 15 days during the treatment period using a MINI-Pam portable modulated chlorophyll fluorometer(Zealquest Scientific Technology Co., Ltd, Shanghai, China). The data were collected from the same seedlings at the same sites on different leaves from the bottom up. The parameters were evaluated using the following formulae.\u003c/p\u003e \u003cp\u003eMaximum photochemical efficiency (Fv/Fm) = (Fm\u0026thinsp;\u0026minus;\u0026thinsp;Fo)/Fm\u003c/p\u003e \u003cp\u003ePotential photochemical efficiency (Fv/Fo) = (Fm\u0026thinsp;\u0026minus;\u0026thinsp;Fo)/Fo\u003c/p\u003e \u003cp\u003eAt the harvest, fresh leaf samples (0.1 g) were cleaned using deionized water to remove any surface contamination. Chlorophyll content was determined using the acetone method. The activities of superoxide dismutase (SOD), peroxisome (POD), and catalase (CAT) were determined using the nitrogen blue tetrazolium, guaiacol, and trace methods, respectively. Malondialdehyde (MDA), proline (Pro), and soluble sugar (SS) contents were determined using the thiobarbituric acid, sulfosalicylic acid, and anthrone colorimetric methods, respectively. Each treatment was repeated three times for each physiological index. The kits for determining SOD, POD, and CAT activities and chlorophyll, MDA, Pro, and SS contents were purchased from Beijing Solaibao Technology Co.(Beijing, China) and used as per the manufacturer\u0026rsquo;s instructions. The levels of hormones [strigolactones (SLs), indole acetic acid (IAA), gibberellic acid (GA), and abscisic acid (ABA)] were measured using enzyme-linked immunosorbent assay (ELISA; EnzymeLink Biologics, Shanghai, China) as per the manufacturer\u0026rsquo;s instructions. The content of each hormone indicator was evaluated using 3 replicates using a microplate reader.\u003c/p\u003e \u003cdiv id=\"Sec10\" class=\"Section2\"\u003e \u003ch2\u003eStatistical analysis\u003c/h2\u003e \u003cp\u003eData analysis was performed using the Statistical Package for the Social Sciences (SPSS 21.0). Potential differences among various treatments were analyzed using Duncan\u0026rsquo;s multiple-comparison tests (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05). All data in the figures and tables are the original data presented as mean\u0026thinsp;\u0026plusmn;\u0026thinsp;standard error (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SE). \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05 was considered significant. The results were plotted using origin 2019 and TB tools.\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003eMycorrhizal properties\u003c/h2\u003e \u003cp\u003eThe AMF colonization rate and intensity and spore number in the uninoculated groups were zero. The colonization rate of AMF-inoculated seedlings in the CK AM was as high as 87.58%. Salt and drought stresses significantly reduced the AMF colonization rate in the seedlings (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05), the rate was slightly higher in the D-AM group than in the S-AM group. AMF colonization intensity was significantly higher in the CK-AM group than in the D-AM group and was significantly higher in the D-AM group than in the S-AM group (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05). The number of spores corresponded to the reproductive strategy of AMF, with a exhibiting a trend of CK-AM\u0026thinsp;\u0026gt;\u0026thinsp;S-AM\u0026thinsp;\u0026gt;\u0026thinsp;D-AM (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05) (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\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\u003eEffect of AMF inoculation in the roots of \u003cem\u003eAlhagi sparsifolia\u003c/em\u003e seedlings under salt and drought stresses\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"5\"\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 \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTreatment\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAMF\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAMF Colonization rate/(%)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eAMF Colonization intensity/(%)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eNumber of AMF spores\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eCK\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0\u0026thinsp;\u0026plusmn;\u0026thinsp;0c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0\u0026thinsp;\u0026plusmn;\u0026thinsp;0d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0\u0026thinsp;\u0026plusmn;\u0026thinsp;0d\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e87.58\u0026thinsp;\u0026plusmn;\u0026thinsp;2.89a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e18.66\u0026thinsp;\u0026plusmn;\u0026thinsp;0.94c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e20.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.58a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0\u0026thinsp;\u0026plusmn;\u0026thinsp;0c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0\u0026thinsp;\u0026plusmn;\u0026thinsp;0d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0\u0026thinsp;\u0026plusmn;\u0026thinsp;0d\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e65.34\u0026thinsp;\u0026plusmn;\u0026thinsp;2.83b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e28.97\u0026thinsp;\u0026plusmn;\u0026thinsp;1.79b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e12.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.58c\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0\u0026thinsp;\u0026plusmn;\u0026thinsp;0c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0\u0026thinsp;\u0026plusmn;\u0026thinsp;0d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0\u0026thinsp;\u0026plusmn;\u0026thinsp;0d\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e62.88\u0026thinsp;\u0026plusmn;\u0026thinsp;2.44b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e48.17\u0026thinsp;\u0026plusmn;\u0026thinsp;1.62a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e13.67\u0026thinsp;\u0026plusmn;\u0026thinsp;0.33b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eCK, D, and S represent blank control, drought stress, and salt stress, respectively. NM and AM represent no inoculation and inoculation with AMF, respectively. The results are expressed as the mean\u0026thinsp;\u0026plusmn;\u0026thinsp;standard error of three replicates. Duncan\u0026rsquo;s test revealed that the parameters differed significantly (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05) among the treatments.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eApparent Seedling Growth\u003c/h3\u003e\n\u003cp\u003eAMF inoculation significantly (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05) promoted the height and basal stem length in \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings, whereas salt and drought stresses had no significant effect on them. Length, surface area, tip number, and specific length of the root were significantly higher in the AM groups than in the NM groups. The root length of seedlings in the S-AM group was significantly lower than that in the D-AM group (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05) (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Under both stress and non-stress conditions, AMF inoculation significantly promoted the growth of above- and belowground parts of \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings; however, the non-stress groups exhibited a stronger promotional effect than the stress groups.\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\u003eEffect of AMF inoculation on the height, basal stem diameter, and root growth of \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings under salt and drought stresses\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"8\"\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 \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTreatment\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAMF\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePlant height growth rate/(%)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eBasal stem growth rate/(%)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eRoot length/(cm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eRoot surface area/(cm\u003csup\u003e2\u003c/sup\u003e)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eNumber of root tips\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eSpecific root length/(m\u0026middot;g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eCK\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.87\u0026thinsp;\u0026plusmn;\u0026thinsp;0.41b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.51\u0026thinsp;\u0026plusmn;\u0026thinsp;0.12c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e13.21\u0026thinsp;\u0026plusmn;\u0026thinsp;0.11d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e3.58\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e6.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.58c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e1.56\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e8.06\u0026thinsp;\u0026plusmn;\u0026thinsp;0.58a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.72\u0026thinsp;\u0026plusmn;\u0026thinsp;0.31a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e64.61\u0026thinsp;\u0026plusmn;\u0026thinsp;0.81a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e24.97\u0026thinsp;\u0026plusmn;\u0026thinsp;1.26a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e37.67\u0026thinsp;\u0026plusmn;\u0026thinsp;3.84a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e3.30\u0026thinsp;\u0026plusmn;\u0026thinsp;0.18a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.56\u0026thinsp;\u0026plusmn;\u0026thinsp;0.19b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.44\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e10.74\u0026thinsp;\u0026plusmn;\u0026thinsp;0.64d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2.68\u0026thinsp;\u0026plusmn;\u0026thinsp;0.19c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e5.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.58c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e1.35\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e7.61\u0026thinsp;\u0026plusmn;\u0026thinsp;0.43a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.36\u0026thinsp;\u0026plusmn;\u0026thinsp;0.14ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e54.78\u0026thinsp;\u0026plusmn;\u0026thinsp;0.69b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e18.21\u0026thinsp;\u0026plusmn;\u0026thinsp;1.32b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e22.00\u0026thinsp;\u0026plusmn;\u0026thinsp;2.08b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e3.58\u0026thinsp;\u0026plusmn;\u0026thinsp;0.20a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.34\u0026thinsp;\u0026plusmn;\u0026thinsp;0.17b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.35\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e9.47\u0026thinsp;\u0026plusmn;\u0026thinsp;0.23d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2.51\u0026thinsp;\u0026plusmn;\u0026thinsp;0.27c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e3.67\u0026thinsp;\u0026plusmn;\u0026thinsp;0.88c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e1.46\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e7.44\u0026thinsp;\u0026plusmn;\u0026thinsp;0.22a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.88\u0026thinsp;\u0026plusmn;\u0026thinsp;0.21bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e44.28\u0026thinsp;\u0026plusmn;\u0026thinsp;0.68c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e17.57\u0026thinsp;\u0026plusmn;\u0026thinsp;1.22b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e20.67\u0026thinsp;\u0026plusmn;\u0026thinsp;3.48b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e3.56\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eCK, D, and S represent blank control, drought stress, and salt stress, respectively. NM and AM represent no inoculation and inoculation with AMF, respectively. The results are expressed as the mean\u0026thinsp;\u0026plusmn;\u0026thinsp;standard error of three replicates. Duncan\u0026rsquo;s test revealed that the parameters differed significantly (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05) between the treatments.\u003c/p\u003e\n\u003ch3\u003eBiomass And Root To Shoot Ratio In The Seedlings\u003c/h3\u003e\n\u003cp\u003eThe fresh and dry weights of above- and belowground parts of \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings increased after AMF inoculation, with the fresh and dry weights of belowground parts increasing significantly (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05) (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Fresh and dry weights were significantly lower in the AM group than in the CK-AM group under salt and drought stresses, indicating that salt and drought stresses had a negative effect on the inoculated AMF. Among them, the dry weight of the belowground part of the S-AM group was significantly lower than that of the D-AM group, indicating that salt stress had a greater effect on the growth of the belowground part of seedlings than drought stress (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05). The root to shoot ratio is an important indicator of plant\u0026rsquo;s allocation of resources. The root to shoot ratios of dry and fresh weights in the AM group were significantly greater than those of the NM group. This indicated that with the assistance of AMF, \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings allocated more resources for the growth and development of the belowground parts. Compared with the CK-AM group, salt and drought stresses significantly promoted an increase in the root to shoot ratio of fresh weight in the AMF-inoculated groups; however, no significant difference existed between the D-AM and S-AM groups.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e\n\u003ch3\u003eN And P Contents\u003c/h3\u003e\n\u003cp\u003eSalt and drought stresses impair the root growth of \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings, thereby reducing the plant\u0026rsquo;s uptake and use of N and P elements. The CK-NM group exhibited higher %N and N content than the D-NM and S-NM groups. Both %N and N content were higher in the CK-AM group than in the D-AM and S-AM groups, with significant difference in their N contents (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05). The P content exhibited the same trend as the N content; however, the variation was greater, with the AM group exhibiting 5\u0026ndash;6-fold higher P content than the NM group. The %P and P content of the D-AM group was significantly higher than that of the S-AM group (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05) (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e), indicating a stronger inhibitory effect of salt stress on access of seedlings to nutrients. Collectively, N and P contents were significantly higher in the no-inoculation group than in the other five groups under salt stress. The N and P contents in the AM group were significantly lower than those in the NM group under salt or drought stress, indicating that AMF inoculation could significantly reduce N and P contents.\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\u003eEffect of AMF inoculation on the N and P contents of \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings under salt and drought stresses\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\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 \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTreatment\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAMF\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eN%\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eNitrogen content/(mg)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eP%\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePhosphorus content/(mg)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eN/P\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eCK\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.79\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.56\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05cd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.13\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.63\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1.58\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08c\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.22\u0026thinsp;\u0026plusmn;\u0026thinsp;0.19a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e6.01\u0026thinsp;\u0026plusmn;\u0026thinsp;0.65a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3.50\u0026thinsp;\u0026plusmn;\u0026thinsp;0.11a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e9.48\u0026thinsp;\u0026plusmn;\u0026thinsp;0.64a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.63\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04d\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.71\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.35\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01cd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.82\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01de\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.13\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2.09\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.01\u0026thinsp;\u0026plusmn;\u0026thinsp;0.13ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3.97\u0026thinsp;\u0026plusmn;\u0026thinsp;0.30b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3.13\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e6.19\u0026thinsp;\u0026plusmn;\u0026thinsp;0.37b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.64\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06d\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.64\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.93\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.53\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.63\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e3.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.91\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3.26\u0026thinsp;\u0026plusmn;\u0026thinsp;0.16bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.54\u0026thinsp;\u0026plusmn;\u0026thinsp;0.23c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e4.35\u0026thinsp;\u0026plusmn;\u0026thinsp;0.52c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.76\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06d\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eCK, D, and S represent blank control, drought stress, and salt stress, respectively. NM and AM represent no inoculation and inoculation with AMF, respectively. The results are expressed as the mean\u0026thinsp;\u0026plusmn;\u0026thinsp;standard error of three replicates. Duncan\u0026rsquo;s test revealed that the parameters differed significantly (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05) between the treatments.\u003c/p\u003e\n\u003ch3\u003eChlorophyll Content And Chlorophyll Fluorescence\u003c/h3\u003e\n\u003cp\u003e \u003c/p\u003e \u003cp\u003eChlorophyll a, chlorophyll b, chlorophyll content, and chlorophyll a/b exhibited following trend under salt or drought stress and AMF inoculation: CK-AM\u0026thinsp;\u0026gt;\u0026thinsp;CK-NM\u0026thinsp;\u0026gt;\u0026thinsp;D-AM\u0026thinsp;\u0026gt;\u0026thinsp;S-AM\u0026thinsp;\u0026gt;\u0026thinsp;D-NM\u0026thinsp;\u0026gt;\u0026thinsp;S-NM. Therefore, AMF promoted the accumulation of chlorophyll in \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings, and the promotional effect was greater under salt stress than under drought stress. Chlorophyll fluorescence is a rapid, nondestructive method for detecting characteristics of plants under stress and photosynthetic performance. The maximum photochemical efficiency (Fv/Fm) and potential photochemical efficiency (Fv/Fo) of the D-NM and S-NM groups exhibited a significant downward trend with increased duration of salt and drought stresses, with the S-NM group exhibiting the fastest and lowest decrease. The Fv/Fm and Fv/Fo values of the CK-NM and CK-AM groups exhibited a stable trend over time. At day 60, the Fv/Fm and Fv/Fo values of the CK-AM group were significantly higher than those of the D-AM and S-AM groups (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05). Comparing the AM and NM groups under salt and drought stresses, respectively, it was clearly observed that AMF inoculation significantly alleviated the effects of salt and drought stresses on the Fv/Fm and Fv/Fo values of the seedlings, particularly in the S-AM group (Fig.\u0026nbsp;2).\u003c/p\u003e \u003cp\u003e \u003cb\u003eFigure\u0026nbsp;2\u003c/b\u003e Effect of AMF inoculation on chlorophyll content and chlorophyll fluorescence in \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings under salt and drought stresses. CK, D, and S represent blank control, drought stress, and salt stress, respectively. NM and AM represent no inoculation and inoculation with AMF, respectively. The results are expressed as the mean\u0026thinsp;\u0026plusmn;\u0026thinsp;standard error of three replicates. Duncan\u0026rsquo;s test revealed that the parameters differed significantly (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05) between the treatments.\u003c/p\u003e \u003cp\u003e \u003cb\u003eActivities of antioxidant enzymes\u003c/b\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eFigure 3 shows that both salt or drought stress and AMF inoculation significantly increased the activities of antioxidant enzymes SOD, POD, and CAT in \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05). The S-AM group exhibited the highest antioxidant enzyme activity. SOD activity was significantly higher in the AM group than in the NM group under salt and drought stresses. It was significantly higher in the S-AM group than in the D-AM group, with no significant difference between AM and NM in the CK group. POD activity was significantly higher in the AM group than in the NM group; it was significantly higher under salt stress than under drought stress and was significantly higher under drought stress than in the blank control. The trend of CAT activity was the same as that of POD. This indicated that AMF inoculation increased the antioxidant enzyme activity in the seedlings in response to the damage caused by salt and drought stresses.\u003c/p\u003e \u003cp\u003e \u003cb\u003eFigure\u0026nbsp;3\u003c/b\u003e Effect of AMF inoculation on the activities of antioxidant enzymes (SOD, POD, and CAT) in \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings under salt and drought stresses. CK, D, and S represent blank control, drought stress, and salt stress, respectively. NM and AM represent no inoculation and inoculation with AMF, respectively. The results are expressed as the mean\u0026thinsp;\u0026plusmn;\u0026thinsp;standard error of three replicates. Duncan\u0026rsquo;s test revealed that the parameters differed significantly (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05) between the treatments.\u003c/p\u003e\n\u003ch3\u003eMda, Pro, And Ss Contents\u003c/h3\u003e\n\u003cp\u003eMDA, Pro, and SS contents increased in \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings under salt and drought stresses (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e4\u003c/span\u003e). The MDA content decreased significantly after AMF inoculation, exhibiting a trend CK-AM\u0026thinsp;\u0026lt;\u0026thinsp;D-AM\u0026thinsp;\u0026lt;\u0026thinsp;S-AM. Pro and SS contents significantly increased after AMF inoculation, and they were significantly higher under salt stress than under drought stress (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05). Therefore, AMF could protect itself from salt and drought stresses to some extent by assisting \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings to accumulate osmoregulatory substances.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e\n\u003ch3\u003eHormonal Levels\u003c/h3\u003e\n\u003cp\u003eSalt and drought stresses significantly increased the IAA and ABA levels, reduced the GA level (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05), and had no significant effect on the SLs level (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.05) in \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e5\u003c/span\u003e). After AMF inoculation, IAA level significantly increased with its level significantly lower in the S-AM group than in the D-AM group; ABA level significantly decreased with its level significantly higher in the S-AM group than in the D-AM group. The GA level was significantly higher in the S-AM group than in the S-NM group but was significantly lower in the D-AM group than in the D-NM group. This indicated that drought and salt stresses regulated the GA level of AMF-inoculated seedlings via various ways. The SLs level was significantly reduced after AMF inoculation, with no significant difference between the salt- and drought-stress groups.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e\n\u003ch3\u003eCorrelation Analysis\u003c/h3\u003e\n\u003cp\u003eThe heat map and Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e show the correlation between various indicators of growth and physiology of \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings under the six treatments, namely, CK-NM, D-NM, S-NM, CK-AM, D-AM, and S-AM (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e6\u003c/span\u003e; Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). The CK-NM, D-NM, and S-NM groups exhibited negative correlations for morphological growth and most physiological indicators and positive correlations for ABA and MDA levels, particularly under salt stress. The positive correlation among growth, biomass accumulation, and chlorophyll content in the above- and belowground parts of seedlings in the CK-AM group was very strong after AMF inoculation. This indicated that AMF inoculation significantly promoted the growth of \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings under non-stress conditions. AMF inoculation alleviated salt- and drought-stress-induced damage to seedlings, which was highlighted by the increased antioxidant enzyme activity and activated osmoregulatory systems of seedlings under salt stress. Antioxidant enzyme activity and osmoregulatory substance content exhibited a significant positive correlation in the S-AM group.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eEffect of salt or drought stress, AMF inoculation, and their interactions on all measured response variables tested using multivariate ANOVA\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\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 \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eindicators\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eDrought\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSalt\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eAMF\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eDrought*AMF\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSalt*AMF\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAMF Colonization rate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e33.279***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e41.037***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1769.429***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e33.279***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e41.037***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAMF Colonization intensity\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e6.032*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e49.421***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e377.183***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e6.032*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e49.421***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eNumber of AMF spores\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e110.979***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e66.083***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1734.289***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e110.979***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e66.083***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePlant height growth rate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.194ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.446ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e49.299***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.007ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.002ns\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBasal stem growth rate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.286ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e6.984*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e26.973***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.719ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e3.449ns\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRoot length\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e12.755**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e17.163**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e101.362***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e9.878**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e9.878**\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRoot surface area\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e5.520*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e21.143**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e354.611***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.976ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e10.043**\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eNumber of root tips\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e17.908**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e21.942**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e461.500***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e10.489**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e12.172**\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSpecific root length\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.146ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.518ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e464.981***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e4.723*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2.469ns\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eFresh weight above ground\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e19.456**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e27.574***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e15.686**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e5.338*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e3.574ns\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eFresh weight below ground\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e9.000*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e10.562**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e108.372***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.174ns\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDry weight above ground\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e8.000*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e18.00**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e7.111*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.500ns\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDry weight below ground\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e19.853**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e74.206***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e250.373***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e10.676**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e19.853**\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRoot to shoot ratio(DW)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.536ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.051ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e56.146***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.737ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.678ns\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRoot to shoot ratio(FW)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.106ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.854ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e177.787***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e17.006**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e10.349**\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e%N\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2.192ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e5.481*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e14.546**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.523ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.683ns\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eN content\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e13.777**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e31.152***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e52.944***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e9.114*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e12.193**\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e%P\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e9.783**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e50.434***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e552.846***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.082ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2.663ns\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eP content\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e26.182***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e68.614***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e261.854***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e14.247**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e31.145***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eN/P\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e24.751***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e246.347***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1477.015***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e22.872***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e174.026***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eChlorophyll a concentration\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e218.337***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e672.622***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e261.534***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e28.105***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e38.587***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eChlorophyll b concentration\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e68.948***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e110.639***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e40.615***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e19.144**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e5.868*\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eChlorophyll contentration\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e18.678**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e164.56***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e81.694***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e77.101***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e77.101***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eChlorophyll a/b ratio\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e249.785***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e649.343***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e248.531***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.412ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e11.139**\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eFv/Fm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e13.135**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e49.662***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.273ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.041ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e4.054ns\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eFv/Fo\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e13.374**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e37.668***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.900ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.308ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e3.115ns\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSuperoxide dismutase SOD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e23.630***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e110.041***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e92.768***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e6.692*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e22.075**\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePeroxidase POD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e16.071**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e79.117***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e26.504***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.294ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.066ns\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCatalase CAT\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e29.390***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e220.195***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e240.587***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.691ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e45.789***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMalondialdehyde MDA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e73.554***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e295.275***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e111.881***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.250ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.324ns\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eProline Pro\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e103.597***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1255.071***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1287.975***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e15.402**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e219.118***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSoluble sugar SS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e5.960*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e26.601***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e63.209***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.136ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e5.467*\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eIndoleacetic acid IAA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.434ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e7.261*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2715.657***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e101.609***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e456.749***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eGibberellin GA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e67.372***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e287.883***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e23.061***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e28.138***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e154.547***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAbscisic acid ABA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e611.830***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1304.326***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e220.300***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e60.476***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e43.074***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eStrigolactones SLs\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2.247ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e9.867**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e50.724***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.044ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.147ns\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"6\"\u003eF-values are followed by \u003cem\u003eP\u003c/em\u003e-values\u003c/td\u003e\u003c/tr\u003e \u003ctr\u003e\u003ctd colspan=\"6\"\u003ens \u003cem\u003eP\u003c/em\u003e ༞0.05\u003c/td\u003e\u003c/tr\u003e \u003ctr\u003e\u003ctd colspan=\"6\"\u003e*\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05\u003c/td\u003e\u003c/tr\u003e \u003ctr\u003e\u003ctd colspan=\"6\"\u003e**\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.01\u003c/td\u003e\u003c/tr\u003e \u003ctr\u003e\u003ctd colspan=\"6\"\u003e***\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003e \u003cem\u003eA. sparsifolia\u003c/em\u003e is an ecologically important plant. The adult plants are ideal for ecological restoration in arid areas. However, salt and drought stresses endanger the morphological establishment of seedlings, which is a key stage in the growth of \u003cem\u003eA. sparsifolia\u003c/em\u003e. Therefore, these stresses limit the process of live regeneration of \u003cem\u003eA. sparsifolia\u003c/em\u003e, and the stability of the population could not be maintained. In this study, we employed the natural environment of the Tarim River basin with a soil salt content of 0.41\u0026ndash;6.28% (Zhong et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) and the salt tolerance range of 0.4\u0026ndash;0.8% for \u003cem\u003eA. sparsifolia\u003c/em\u003e seed germination (Zhang et al. 2009). Based on this condition and our previous study (Zhang et al. 2016; Zainur 2021; GAO et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), we conducted this study under salt stress (0.6%) and drought stress (30% \u0026plusmn; 5%) to verify whether AMF can improve the salt and drought tolerance of \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings and to reveal the survival strategies during the seedling period by altering apparent growth, photosynthetic efficiency, antioxidant enzyme activity, and levels of osmoregulatory substances and hormones.\u003c/p\u003e\n\u003ch3\u003eEffects Of Amf Inoculation On Salt- Or Drought-induced Damage In The Seedlings\u003c/h3\u003e\n\u003cp\u003eStudies have reported that the root system of \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings can establish a good symbiosis with AMF, which is a highly clumped mycorrhizal-dependent plant (Wang et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). The rate and intensity of AMF colonization were significantly reduced under salt and drought stresses, indicating that salt and drought stresses significantly inhibited the growth of AMF. The present study reported that although the colonization intensity of AMF under salt stress was significantly lower than that under drought stress, the spore number of AMF was significantly higher under salt stress than under drought stress. This suggested that the massive accumulation of AMF spores may be an effective reproductive strategy for its successful survival in adverse conditions.\u003c/p\u003e \u003cp\u003eSalt and drought stresses inhibited the growth and live regeneration processes in the seedlings (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e); however, AMF promoted host plant growth by expanding the total root uptake area and improving uptake of nutrients by the plant (Diagne et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The height and basal stem length of \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings significantly increased after AMF inoculation under both stress and non-stress conditions. After AMF inoculation, the root length increased by 3.8, 4.1, and 3.6-fold in the non-stressed, drought-stressed, and salt-stressed groups, respectively. The specific root length and root to shoot ratio increased by 1 fold in the D-AM groups, indicating that AMF responded to the damage due to drought stress by enhancing the root growth of the seedlings. Adaptation in the form of changes in root morphology and microbial cooperation require the allocation of carbon from photosynthesis to compete with the belowground carbon sinks, either to promote root growth or to transfer to symbiotic partners (Cao et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Plant biomass allocation is the result of a combination of biotic and abiotic factors, and the exponential increase in the root to shoot ratio of the seedlings under abiotic and non-stress factors confirmed the significant promotion of root growth by the biotic factor AMF. The exponential increase in the specific root length further confirmed the beneficial effect of AMF on the increase of root length during the root growth of the seedlings.\u003c/p\u003e\n\u003ch3\u003eEffects Of Amf Inoculation On N And P Contents Under Salt Or Drought Stress\u003c/h3\u003e\n\u003cp\u003eN and P are the two elements that are essential for plant growth. Notably, AMF inoculation significantly promoted the uptake of elemental P. The low fixation of P in soil and its rapid precipitation with soil minerals result in low effective P content in soil, whereas AMF contributes to 80% of total P uptake and has the potential to secrete acid phosphatase and organic acids that directly promote P uptake (Masao et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Song et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). The importance of AMF in P uptake was demonstrated by the 4\u0026ndash;5-fold increase in the P uptake after AMF inoculation in this study. The contribution of AMF to N uptake is usually smaller than that to P uptake by plants; however, AMF can still transfer N to the host (Wang et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). In this study, the N content in the AMF-inoculated groups was higher than that in the non-inoculated groups and was more influenced by stress factors. The AMF-inoculated seedlings had the highest total N content under non-stress conditions. The AMF-inoculated seedlings had a higher total N content than the non-inoculated seedlings under salt and drought stresses, and the non -inoculated seedlings had the lowest total N content under salt stress. Both salt and drought stresses and AMF inoculation exhibited important effects on the growth of seedlings. N:P is an accurate indicator of the nutrient limitation for plant growth. N:P of \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings obtained in this study ranged from 0.6 to 3.1; this indicated that AMF helped the seedlings in increasing their P content while leaving them relatively deficient in N, a phenomenon that was more pronounced under salt and drought stresses.\u003c/p\u003e\n\u003ch3\u003eEffects Of Amf Inoculation On The Photosynthesis Under Salt Or Drought Stress\u003c/h3\u003e\n\u003cp\u003eChlorophyll content is an indicator of photosynthetic status, developmental stage, and degree of stress (Li et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The chlorophyll content in \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings under salt stress was significantly lower than that under drought stress and was significantly lower in the stress groups than in the non-stress groups. This indicated that both salt and drought stresses can impair the growth of seedlings by reducing their chlorophyll content, and in turn photosynthetic capacity; however, this effect was greater under salt stress. After AMF inoculation, the chlorophyll content of seedlings significantly increased, indicating that the presence of AMF and salt stress had less effect on chlorophyll synthesis. This could be attributed to the substances secreted by AMF, such as cytokinins, as these substances facilitate chloroplast development and increase chlorophyll levels (Wang et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Photosystem II (PSII) is the main light-driven reaction involved in photosynthesis (Wu et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Salt and drought stresses reduced the photochemical efficiency and electron transfer in the PSII reaction center, resulting in lower Fv/Fm and Fv/Fo values than normal in the leaves of \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings. However, Fv/Fm and Fv/Fo values did not significantly decrease with increasing duration of stress after AMF inoculation. This indicated that salt and drought stresses had a negative impact on the photosynthetic physiology of the seedlings, and salt stress had a greater impact; AMF inoculation had a sustained but not significant effect in alleviating the effects of the stresses.\u003c/p\u003e\n\u003ch3\u003eEffects Of Amf Inoculation On The Antioxidant Enzyme Activities Under Salt Or Drought Stress\u003c/h3\u003e\n\u003cp\u003eAMF promotes plant growth, increases the above- and belowground biomass, and mitigates the negative effects of abiotic stresses (Asma et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Dorot\u0026eacute;ia et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Wang et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). AMF inoculation activated the antioxidant enzyme system of \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings and increased the activity of SOD, POD, and CAT, which was particularly significant under salt stress compared with that under drought stress. The antioxidant enzyme activity was most correlated with AMF inoculation in terms of physiological indicators in the seedlings. This could be because of following reasons. First, salt stress increased the accumulation of toxic reactive oxygen species (ROS) and stimulated the seedlings\u0026rsquo; own antioxidant defense system to produce numerous antioxidant enzymes. Second, AMF protected the seedlings from ROS by enhancing antioxidant enzyme activity to maintain an effective antioxidant enzyme system. Furthermore, numerous studies have reported that the antioxidant enzyme activity of AMF-inoculated plants is higher than that of non-inoculated plants in this species (Wang et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The antioxidant enzyme activity was increased under drought stress as well; however, the effect was lower than that under salt stress, presumably because the levels of osmotic pressure and osmoregulatory substances in the seedlings were greater under salt stress than under drought stress.\u003c/p\u003e\n\u003ch3\u003eEffects Of Amf Inoculation On Mda, Pro, And Ss Contents Under Salt Or Drought Stress\u003c/h3\u003e\n\u003cp\u003eSalt stress can directly lead to osmotic stress, primary ion toxicity, and secondary stresses such as nutrient deficiency and oxidative stress in plants, which ultimately hinder plant growth (Yang et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). The effects of drought stress are similar to those of salt stress. Both salt and drought stresses disrupt cell structure and some key physiological functions in plants, e.g., cell membrane damage, reduced photosynthetic capacity, production of ROS, and accumulation of harmful substances (Li et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The results of this study demonstrated that the content of osmoregulatory substances increased in \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings, and the increase was higher under salt stress. Pro is the main store of energy and N under salt stress (Wang et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). After AMF inoculation, the Pro content increased significantly under salt stress. Therefore, AMF helped the plant in accumulating more Pro and overcoming the salt-stress-induced osmotic imbalance.\u003c/p\u003e\n\u003ch3\u003eEffects Of Amf Inoculation On Hormonal Levels Under Salt Or Drought Stress\u003c/h3\u003e\n\u003cp\u003eAMF not only improves macroscopic plant growth and N and P uptake but also regulates microscopic intracellular hormonal levels. AMF colonization relies on the release of SLs into the root zone to germinate spores and attract mycelium to the root system (Ma et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In this study, the level of SLs in the AMF-inoculated groups was significantly lower than that in the non-inoculated groups. It was assumed that SLs were released into the soil for the colonization of AMF with the root system in the AMF-inoculated group. SLs can regulate the process of adaptation of plants to adverse conditions (Liao et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). In this study, the level of SLs was significantly lower in the non-inoculated group under salt stress than in the blank control group. Compared with the blank control group, the increase in the IAA and ABA levels and decrease in the GA level under salt and drought stresses demonstrated the self-regulation ability of the seedlings under salt and drought stresses. The significant increase in the IAA level and significant decrease in the GA level in this study supported the idea that significant changes in hormonal levels are also a survival strategy for seedlings to mitigate the effects of salt and drought stresses with the help of AMF (Huang et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). AMF enhanced the tolerance of the seedlings to salt and drought stresses through a combination of mechanisms, reaffirming the critical role of mycorrhizal fungi as a major component of terrestrial ecosystems, particularly in the restoration of degraded desert ecosystems (Meng et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2008\u003c/span\u003e).\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eIn summary, we can conclude that \u003cem\u003eA. sparsifolia\u003c/em\u003e\u0026ndash;AMF symbiosis mitigated the negative effects of salt and drought stresses on \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings by altering plant growth and physiology. The mechanisms are as follows. AMF inoculation promoted the growth of above- and belowground parts, facilitating more nutrient gain and biomass accumulation. It better balanced light energy distribution in photochemical and non-photochemical processes. It increased antioxidant enzyme activity to scavenge and reduce the levels of ROS. It increased the levels of soluble substances to improve osmoregulation. It up- or downregulated hormonal levels to induce hormone signaling responses, ultimately resulting in better salt and drought tolerance by the seedlings. Therefore, AMF has important potential application in the regeneration of \u003cem\u003eA. sparsifolia\u003c/em\u003e and restoration of degraded desert ecosystems in arid zones.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgments\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis work was financially supported by the National Natural Science Foundation of China (No. 42067067) and the Natural Science Foundation of Xinjiang Uygur Autonomous Region (No.2020D01A74). Thanks to Dr. Jingbo Zhang for guidance in TB tools data processing software.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor Contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eFormal analysis: Xiaodong Ma,Aili Yilinuer.\u003c/p\u003e\n\u003cp\u003eResources:Dawuti Maigepiretiguli.\u003c/p\u003e\n\u003cp\u003eSoftware: Haiou Wang.\u003c/p\u003e\n\u003cp\u003eSupervision: Xiaonan Chen.\u003c/p\u003e\n\u003cp\u003eWriting - original draft: Xiaonan Chen.\u003c/p\u003e\n\u003cp\u003eWriting - review: Xiaodong Ma.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDeclarations\u003c/strong\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eConflict of interest The author declared that they have no conflict of interest.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAbdelmalik AM, Alsharani TS, Al-Qarawi AA, Ahmed A, Aref IM (2020) Response of growth and drought tolerance of Acacia seyal Del. seedlings to arbuscular mycorrhizal fungi[J]. 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Res Soil Water Conserv 25(04):134\u0026ndash;138. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.13869/j.cnki.rswc.2018.04.020\u003c/span\u003e\u003cspan address=\"10.13869/j.cnki.rswc.2018.04.020\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\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":true,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"plant-growth-regulation","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"grow","sideBox":"Learn more about [Plant Growth Regulation](https://www.springer.com/journal/10725)","snPcode":"10725","submissionUrl":"https://submission.nature.com/new-submission/10725/3","title":"Plant Growth Regulation","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"The Tarim River Basin, Alhagi sparsifolia seedlings, Plant–Microbe Interaction, Antioxidant enzyme system, Phytohormones","lastPublishedDoi":"10.21203/rs.3.rs-2374175/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-2374175/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003e \u003cem\u003eAlhagi sparsifolia\u003c/em\u003e is a dominant species in the lower Tarim River desert ecosystem and an important mycorrhizal symbiont. However, it is unclear how its seedlings cope with salt and drought stresses and develop effective survival strategies with the assistance of arbuscular mycorrhizal fungi (AMF) during the vulnerable growth stage. Therefore, in this study, we investigated the effects of salt or drought stress on the growth rate; nutrient uptake; photosynthesis; and the levels of antioxidant enzymes, osmoregulatory substances, and hormones in \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings. We included following six groups: control/CK-NM (without salt or drought stress and no AMF inoculation), D-NM (drought stress and no AMF inoculation), S-NM (salt stress and no AMF inoculation), CK-AM (AMF inoculation and no salt or drought stress), D-AM (drought stress and AMF inoculation), and S-AM (salt stress and AMF inoculation). The results revealed that AMF inoculation promoted seedling growth, particularly root growth and phosphorus nutrient uptake, in \u003cem\u003eA. sparsifolia\u003c/em\u003e. Salt and drought stresses negatively affected the growth, photosynthetic capacity, and nutrient accumulation in the above- and below-ground parts of the seedlings and stimulated the antioxidant defense system and accumulation of osmoregulatory substances in them. AMF inoculation under salt and drought stresses could alleviate toxic symptoms in \u003cem\u003eA. sparsifolia\u003c/em\u003e by promoting root growth, enhancing nutrient uptake, activating antioxidant enzyme activity, and regulating hormonal levels. These effects of AMF were mainly reflected in root growth under drought stress and antioxidant enzyme activity under salt stress. The beneficial effect of AMF under salt stress was better than that under drought stress. This study demonstrated that AMF plays a significant role in assisting \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings to quickly pass through the vulnerable growth stage under salt and drought stresses. Therefore, \u003cem\u003eA. sparsifolia\u003c/em\u003e seedlings with AMF have potential application in restoration of desert ecosystem.\u003c/p\u003e","manuscriptTitle":"Mycorrhizal fungal colonization promotes apparent growth and physiology of Alhagi sparsifolia seedlings under salt or drought stress at vulnerable developmental stage","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2022-12-20 19:22:27","doi":"10.21203/rs.3.rs-2374175/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Major revisions","date":"2023-01-20T03:17:11+00:00","index":"","fulltext":""},{"type":"reviewerAgreed","content":"","date":"2022-12-17T00:07:55+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2022-12-16T14:20:29+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"Plant Growth Regulation","date":"2022-12-16T06:49:02+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2022-12-14T11:12:15+00:00","index":"","fulltext":""},{"type":"submitted","content":"Plant Growth Regulation","date":"2022-12-13T08:31:47+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"plant-growth-regulation","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"grow","sideBox":"Learn more about [Plant Growth Regulation](https://www.springer.com/journal/10725)","snPcode":"10725","submissionUrl":"https://submission.nature.com/new-submission/10725/3","title":"Plant Growth Regulation","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"fab6e874-c593-4ee5-b57d-5cc534d5f57e","owner":[],"postedDate":"December 20th, 2022","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2023-10-16T20:25:36+00:00","versionOfRecord":{"articleIdentity":"rs-2374175","link":"https://doi.org/10.1007/s10725-023-00996-0","journal":{"identity":"plant-growth-regulation","isVorOnly":false,"title":"Plant Growth Regulation"},"publishedOn":"2023-04-05 20:22:47","publishedOnDateReadable":"April 5th, 2023"},"versionCreatedAt":"2022-12-20 19:22:27","video":"","vorDoi":"10.1007/s10725-023-00996-0","vorDoiUrl":"https://doi.org/10.1007/s10725-023-00996-0","workflowStages":[]},"version":"v1","identity":"rs-2374175","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-2374175","identity":"rs-2374175","version":["v1"]},"buildId":"7rjqhiLT3MXkJMwkYKINL","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}