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Article

Diversity and Community Structure of Rhizosphere Arbuscular Mycorrhizal Fungi in Songnen Grassland Saline–Alkali-Tolerant Plants: Roles of Environmental Salinity and Plant Species Identity

1
College of Animal Science and Technology, Northeast Agricultural University, Harbin 150030, China
2
State Key Laboratory of Ecological Safety and Sustainable Development in Arid Lands, Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences, Urumqi 830011, China
*
Author to whom correspondence should be addressed.
Agronomy 2025, 15(9), 2070; https://doi.org/10.3390/agronomy15092070
Submission received: 24 July 2025 / Revised: 15 August 2025 / Accepted: 27 August 2025 / Published: 28 August 2025
(This article belongs to the Section Agricultural Biosystem and Biological Engineering)

Abstract

The Songnen Grassland, a typical saline–alkali ecosystem in Northeast China, is increasingly degraded by soil salinization. Arbuscular mycorrhizal fungi (AMF) are critical for enhancing plant tolerance to saline–alkali stress via root symbiosis. To investigate the species diversity and community structure of AMF in the rhizosphere of salt-tolerant plants in the Songnen Grassland, this study combined morphological identification with high-throughput sequencing (based on virtual taxa, VTs, from the MaarjAM database) to analyze the composition and distribution characteristics of AMF in the rhizosphere of eight salt-tolerant plant species, including Arundinella anomala, Leymus chinensis, Taraxacum mongolicum and others. Morphological identification revealed a total of 22 AMF species belonging to 7 genera. Among these, the genus Glomus was the dominant genus, comprising eight species (accounting for 36.4% of the total species), followed by the genus Acaulospora (five species, 22.7%), the genus Rhizophagus (four species, 18.2%), the genus Ambispora (two species, 9.1%), and the remaining genera each represented by one species (4.5%). High-throughput sequencing analysis identified a total of 40 virtual taxa (VTs) with clear taxonomic assignments belonging to six genera. The genus Glomus accounted for the highest proportion (34 VTs, 85%) with a relative abundance of 89.33%, representing the overwhelmingly dominant group. Rhizosphere soil electrical conductivity (EC) of the eight plant species indicated a significant gradient (high EC group: A–D and G, 2.07–2.61 mS/cm; low EC group: E, F, H, 0.20–0.48 mS/cm). The AMF diversity in the high EC group was significantly higher than that in the low EC group, indicating that AMF in the rhizosphere of salt-tolerant plants enhanced plant tolerance to high-salt environments, and their diversity did not decrease with increasing salinity but instead remained at a high level. Plant-specific AMF community characteristics were evident. Hierarchical clustering analysis further confirmed that the AMF community composition in the rhizosphere of Taraxacum mongolicum and Vicia amoena differed significantly from that of the other plant species, indicating that plant species have a key driving role in AMF community structure. These findings provide critical insights into the plant–AMF symbiotic mechanisms underlying saline–alkali adaptation and offer a theoretical basis for selecting efficient AMF strains to support ecological restoration of saline–alkali lands.

1. Introduction

The Songnen Grassland is a key component of northeastern China’s terrestrial ecosystems and one of the world’s three major soda saline–alkaline soil hubs. Severe soil salinization threatens regional ecological stability and agricultural productivity [1]. Dominant salts include soda salts (sodium carbonate, sodium bicarbonate), sulfates, and chlorides [2,3,4]. Natural factors (e.g., climatic aridification, parent material weathering) and anthropogenic activities (e.g., large-scale land reclamation, inefficient water use) jointly accelerate soil degradation [5]. Degradation manifests as compaction, lost aggregate structure, and nutrient depletion [6]. These changes directly inhibit plant growth via osmotic stress (reduced water uptake) and ion toxicity (excess Na+ disrupting intracellular Na+/K+ homeostasis) [7]. They also cause nutrient imbalances, constraining sustainable agriculture in the region. To address these challenges, AMF—known for enhancing plant salt tolerance [8], improving soil health [9], and facilitating nutrient uptake—have emerged as a viable biological strategy for saline–alkaline land restoration [10]. AMF are now a critical research priority for mitigating soil salinization in the Songnen Grassland.
AMF are indispensable to terrestrial ecosystems, comprising 5–50% of soil microbial communities [11]. They form symbioses with >80% of terrestrial plants, boosting phosphorus uptake and soil nutrient cycling [12]. AMF also enhance plant pest/disease resistance via root exudates [13] and improve legume nitrogen fixation efficiency [14]. Additionally, their symbiotic structures (e.g., arbuscules, hyphae) and plant root exudates secrete main enzymes such as phosphatases, ureases, and sucrases that stimulate soil microbial activity, optimizing conditions for plant growth [15]. This makes AMF a critical link between plant and soil health [16].
Soil salinization, a global environmental issue, inhibits AMF growth and reproduction [17]. High-salt environments increase soil osmotic pressure, restricting water and nutrient uptake by hyphal cells [18]. High salinity increases soil osmotic pressure, limiting hyphal water and nutrient uptake [19]. Excess Na+ and Cl cause ion toxicity, disrupting intracellular homeostasis and metabolism [20]. Elevated salinity further impairs spore germination by reducing soil suitability [21]. Moreover, salt-tolerant bacteria intensify resource competition, indirectly suppressing germination [22]. These mechanisms collectively weaken AMF adaptability in saline–alkaline environments.
Saline–alkali stress also alters plant community structure, growth, and physiology, weakening plant–AMF symbiosis [23]. AMF intensify plant interspecific competition and participate in population self-thinning under varying conditions [24]. In saline–alkaline soils, plants allocate more energy to osmotic balance and oxidative stress [25], reducing carbon supply to AMF [26]. Since AMF rely entirely on host photosynthates, carbon limitation restricts their reproduction [27]. Salt–alkali stress may also alter root exudate composition, indirectly affecting AMF dynamics [28]. These changes ultimately reduce the fitness of the plant–AMF symbiotic system.
Despite growing research on AMF in saline–alkaline ecosystems [29,30], critical gaps remain. Most studies focus on single plant species or broad environmental factors (e.g., total salt concentration) [31,32], but few have explored how plant species identity (e.g., differences in root exudates, salt tolerance strategies) interacts with specific soda saline–alkaline properties (e.g., high HCO3 concentration, extreme pH) to shape AMF community structure. Additionally, the functional significance of AMF diversity in supporting salt-tolerant plant communities in soda saline–alkaline grasslands—such as the Songnen—has not been fully elucidated. These gaps limit our ability to design targeted AMF-based strategies for restoring degraded saline–alkaline ecosystems.
Terrestrial ecosystem functioning and stability depend on biodiversity, rooted in mechanisms enabling species coexistence. AMF regulate plant competitive dynamics and expand niche space via resource partitioning. They play a critical role in host plant niche construction and community maintenance [33]. This study investigates AMF diversity, community composition, and biotic/abiotic drivers in the rhizospheres of 8 salt-tolerant plants in the Songnen Grassland (a typical soda saline–alkaline ecosystem). We propose the following hypotheses: (1) Higher AMF diversity in the rhizosphere of salt-tolerant plants in the Songnen grassland, and certain salt-tolerant genera (e.g., Glomus) may increase in relative dominance due to their enhanced adaptability. (2) Plant species are the main biotic drivers shaping the community structure of AMF. Testing these hypotheses will deepen our understanding of plant–AMF symbiotic mechanisms under salt stress and provide a theoretical basis for selecting AMF strains for saline–alkaline ecosystem restoration.

2. Materials and Methods

2.1. Study Site and Sample Collection

The research site is situated in the Songnen Grassland, located in Zhaodong City, Heilongjiang Province, China (elevation: 139–140 m; coordinates: 46°2′45″–46°3′12″ N, 125°53′51″–125°54′1″ E; Figure S1). This region has a temperate continental climate, with warm and humid summers (20–24 °C) and cold, dry winters (−18 to −15 °C). The annual average precipitation is approximately 569.1 mm, of which more than 60% occurs between June and September. The soil salinity is dominated by sodium carbonate (Na2CO3) and sodium bicarbonate (NaHCO3), which together account for over 50% of the total salt content and significantly influence the vegetation distribution pattern. Currently, the Songnen grassland ecosystem is facing multiple pressures, including salinization and overgrazing, resulting in noticeable changes in its plant community structure. The main dominant species include Puccinellia tenuiflora (Griseb.) and Leymus chinensis (Trin.).
Against this ecological backdrop, this study investigated eight representative plant species, specifically: Arundinella anomala (No. A, 46°2′46″ N, 125°53′51″ E), Leymus chinensis (No. B, 46°2′46″ N, 125°53′53″ E), Taraxacum mongolicum (No. C, 46°3′10″ N, 125°53′59″ E), Puccinellia tenuiflora (No. D, 46°3′9″ N, 125°53′58″ E), Artemisia mongolica (No. E, 46°3′12″ N, 125°54′1″ E), Artemisia anethifolia (No. F, 46°3′12″ N, 125°54′0″ E), Clematis hexapetala (No. G, 46°2′54″ N, 125°53′51″ E), Vicia amoena (No. H, 46°3′8″ N, 125°54′0″ E). These plants play a significant role in adapting to saline–alkali environments and maintaining the stability of grassland ecosystems, while also reflecting the unique response mechanisms of plant communities in this region to abiotic stress.
Sampled from 8 single-species-dominated patches. Soil electrical conductivity (EC) was measured first, and patches were categorized into high EC (2.07–2.61 mS/cm) and low EC (0.20–0.48 mS/cm) groups before plant sampling. This patch structure ensured homogeneous microenvironments for each sample, minimizing confounding factors from mixed vegetation. Spatial independence was maintained by a 50–200 m distance between adjacent patches and at least 10 m between three healthy individuals selected per species within a patch, reducing cross-contamination of rhizosphere microbial communities and avoiding spatial autocorrelation. Root morphology varied significantly among species: fibrous roots (e.g., Arundinella anomala, Leymus chinensis) provided large surface areas for AMF colonization, taproots (e.g., Taraxacum mongolicum, Vicia amoena) altered rhizosphere microhabitats (e.g., nutrient distribution, oxygen availability), and shallow spreading roots (e.g., Puccinellia tenuiflora) restricted colonization to topsoil—differences that may influence AMF diversity by modifying root–fungal interactions and carbon source (root exudate) availability. Life cycle consistency was critical: All plants were sampled at the late vegetative or early flowering stage (height: 20–50 cm; similar growth vigor), a period of active root growth and AMF association, avoiding biases from seedling (low AMF colonization) or senescent (declining AMF activity) stages. On 17 September 2021, rhizosphere soil (5–20 cm depth, tightly adhering to roots) was collected via multi-point composite sampling; each sample was split for air-drying (morphological analysis) or stored at −80 °C (high-throughput sequencing).

2.2. Laboratory Analyses

AMF spores were isolated from 100 g of air-dried rhizosphere soil using a wet sieving–sucrose centrifugation method [34] with the following: Wet sieving: Soil samples were suspended in deionized water and passed through a series of sieves (250 μm, 106 μm, 45 μm) to retain spores of different sizes. Sucrose centrifugation: The retained material from the 45 μm sieve was transferred to a 50 mL centrifuge tube, mixed with a 50% (w/v) sucrose solution, and centrifuged at 1500 rpm for 5 min. The supernatant (containing spores) was poured onto a 45 μm sieve and rinsed with deionized water to remove sucrose. The purified spore suspension was transferred to a sterile Petri dish and observed under a dissecting microscope (Olympus SZX16, Olympus Corporation, Tokyo, Japan) at 40× magnification. Individual spores were picked using a sterile capillary tube and mounted on glass slides with lactophenol cotton blue stain (0.05% w/v). Slides were sealed with nail polish and incubated at room temperature for 24 h to allow thorough staining. Spores were mounted on microscope slides for morphological observation (shape, color, wall structure, hyphal attachment). Species identification was based on the AMF in China monograph, the International Culture Collection of AMF (INVAM) classification criteria, and the published literature [35]. Spore density (number of spores per 100 g dry soil) and hyphal density (mg/g dry soil) were measured following the method of Mei et al. [36]. Soil electrical conductivity (EC) was measured with a conductivity meter (DDS-307A, Shanghai Leici Instrument Factory, Shanghai, China) and soil pH was determined with a pH meter (PHS-3C, Shanghai Leici Instrument Factory, Shanghai, China), both in 1:5 (soil:water) suspensions [37].
Genomic DNA was extracted from 0.5 g of frozen rhizosphere soil using the PowerSoil DNA Isolation Kit (Mo Bio Laboratories, Carlsbad, CA, USA) following the manufacturer’s instructions. Three biological replicates were performed per plant species. DNA quality was assessed via: (1) 1% agarose gel electrophoresis (integrity, no degradation); (2) NanoDrop (OD260/OD280 = 1.8–2.0, OD260/OD230 > 1.5, purity).
Nested PCR was utilized to amplify the 18S rRNA gene fragment of AMF. The first round of PCR used the universal fungal primers AML1F (5′-ATCAACTTTCGATGGTAGGATAGA-3′) and AML2R (5′-GAACCCAAACACTTTGGTTTCC-3′). The second round (nested PCR) used AMF-specific primers AMV4.5NF (5′-AAGCTCGTAGTTGAATTTCG-3′) and AMDGR (5′-CCCAACTATCCCTATTAATCAT-3′). PCR reactions (25 μL) contained 12.5 μL 2× Taq PCR Master Mix, 1 μL each primer (10 μM), 2 μL DNA template, and 8.5 μL ddH2O. Cycling conditions for the first round: 94 °C for 5 min; 30 cycles of 94 °C for 30 s, 55 °C for 30 s, 72 °C for 45 s; 72 °C for 10 min. Nested PCR conditions: 94 °C for 5 min; 35 cycles of 94 °C for 30 s, 58 °C for 30 s, 72 °C for 45 s; 72 °C for 10 min.
Amplified products were purified using the AxyPrep DNA Gel Extraction Kit (Axygen, Union City, CA, USA) and used to construct Illumina MiSeq libraries. Sequencing was performed on the Illumina MiSeq platform (2 × 300 bp paired-end) by Majorbio Bio-Pharm Technology Co., Ltd. (Shanghai, China).
Raw sequences underwent quality filtering via Trimmomatic (version 0.39) to discard low-quality reads (Q < 20) and adapter sequences. Paired-end reads were merged with FLASH (version 1.2.11), followed by chimeric sequence removal using UCHIME (version 4.2). Non-redundant sequences were then clustered into operational taxonomic units (OTUs) at 97% sequence similarity via UPARSE (version 7.1). Singleton OTUs (those with only one sequence) were excluded to minimize noise.
The most abundant sequence in each OTU was selected as the representative sequence. Taxonomic annotation was performed using the Silva database (version 138) for 18S rRNA gene sequences, with additional validation using the MaarjAM database. OTUs classified as non-AMF (e.g., other fungi, bacteria) were excluded from subsequent analysis.
To analyze the AMF community in the rhizosphere of different plants, we performed high-throughput sequencing and annotated species by comparing sequences to the MaarjAM database. This yielded a total of 40 virtual taxa (VTs) with clear taxonomic assignments, spanning 6 genera. Glomus was the most dominant genus, accounting for 34 VTs (85% of the total). The remaining VTs were distributed among Archaeospora (2 VTs), Diversispora (1 VT), Gigaspora (1 VT), and Paraglomus (2 VTs). Some species detected in morphological analyses were not detected in high-throughput sequencing—likely due to either primer bias (the AMV4.5NF/AMDGR primers used have low affinity for partial genus 18S rRNA genes) or low abundance (below the sequencing detection limit. The MaarjAM database (https://maarjam.ut.ee/; accessed on 7 November 2021), established by Davison et al. [38], compiles all published AM fungal sequences targeting the NS31-AM1 primer region. Using a 97% sequence similarity threshold, the authors defined 356 molecular species and classified each into genus or species levels by aligning with reference DNA sequences from spore-derived taxa. These classifications are termed “virtual taxa (VTs)” [38].

2.3. Statistical Analysis

All data were organized in Excel 2019 (Microsoft Corporation, Redmond, WA, USA) and analyzed using IBM SPSS Statistics 27 (IBM Corp., Armonk, NY, USA). AMF community diversity and richness were quantified using four indices: Sobs: Actual number of distinct AMF species observed per sample (observed richness); Chao1 index: Estimated total species richness accounting for undetected rare species, calculated as:
Chao 1 = S obs + F 1 2 2 F 2
where S obs = observed species number, F 1 = number of singleton species (found in one sample), F 2 = number of doubleton species (found in two samples).
Shannon index ( H ): Combines species richness and evenness, calculated as:
H = i = 1 S p i l n p i
Simpson index ( D ): Emphasizes dominant species, calculated as:
D = 1 i = 1 S p i 2
where S = total species number, p i = proportion of sequences belonging to species i .
Prior to hypothesis testing, all datasets—including diversity indices (Shannon, Simpson, Sobs, Chao1), spore density, and hyphal density—were rigorously assessed for normality using the Shapiro–Wilk test (α = 0.05) and homogeneity of variance via Levene’s test (α = 0.05). Variance stabilization was implemented where required: Shannon and Simpson indices (typically log-normally distributed) underwent natural log transformation (ln[x + 1]), while Sobs and Chao1 (inherently discrete count data) remained untransformed. For datasets satisfying parametric assumptions after transformation, differences among plant species were evaluated using one-way ANOVA followed by Tukey’s HSD post hoc test. Non-parametric datasets were analyzed with the Kruskal–Wallis H test and Dunn’s post hoc correction. Statistical significance was defined at α = 0.05, and all results are reported as mean ± standard error (SE).

3. Results

3.1. AMF Species in the Rhizosphere of Eight Salt-Tolerant Plants

Through morphological identification of rhizosphere soil samples from different plants in the Songnen Grassland, we isolated 22 AMF species belonging to 7 genera. Table 1 summarizes the taxonomic diversity of these AMF, detailing the distribution of species across genera and their relative proportions of the total identified species.
Among the genera, Glomus was the most dominant, comprising eight species (36.4% of the total), followed by Acaulospora with five species (22.7%), Rhizophagus with four species (18.2%), and Ambispora with two species (9.1%). The remaining genera—Claroideoglomus, Gigaspora, and Funneliformis—each contributed one species, accounting for 4.5% of the total species, respectively (Table 1).
Table 2 summarizes the virtual taxa (VTX) of AM fungi isolated from the rhizospheres of different plants via high-throughput sequencing, categorizing these taxa by genus to illustrate taxonomic diversity across groups. At the genus level, Glomus dominated the community with a relative abundance of 89.33% of total sequences. The second most abundant group was Unclassified_class_Glomeromycetes (10.22%), followed by Paraglomus (0.06%), and other genera (collectively 0.06%; Figure 1). At the species level, Glomus Wirsel OTU6 (VTX00202) was the dominant species, contributing 29.27% of the total sequences. This was followed by unclassified_g_Glomus_family_Glomeraceae (uncategorized species, 20.63%). In contrast, Glomus sp. (VTX00304) had the lowest relative abundance (0.05%). These results collectively reflect the key characteristics of microbial distribution in this rhizosphere environment—with Glomus as the dominant genus and distinct species-level variations in abundance (Table 2; Figure 1).

3.2. Distribution of Arbuscular Mycorrhizal Fungi in the Rhizosphere of Eight Salt-Tolerant and Alkali-Resistant Plant Species

There were significant differences in the AMF community structures among the rhizospheres of eight plant species in the Songnen grassland (Table 3). Among the eight soil samples collected, Leymus chinensis, Taraxacum mongolicum, and Puccinellia tenuiflora exhibited the highest AMF diversity, with 15, 13, and 12 species identified, respectively, accounting for 68.2%, 59.1%, and 54.5% of the total AMF species identified via morphological analysis. In contrast, Clematis hexapetala had the lowest AMF diversity, with only seven species isolated, representing 31.8%. Glomus mosseae and Glomus etunicatum were broadly distributed across all samples and emerged as the most prevalent AMF species in the region. Glomus convolutum, Acaulospora laevis, and Rhizophagus intraradices were detected in five samples each. Glomus deserticola was only detected in the rhizosphere soil of Taraxacum mongolicum, suggesting it may be a host-specific or occasional AMF species associated with the rhizosphere of Taraxacum mongolicum.

3.3. Relationship Between AMF and Plant Species

At the species level, Glomus Wirsel OTU6 VTX00202 exhibited the highest relative abundance, accounting for 29.27% of the total sequences, and was the dominant species in the rhizospheres of the sampled plants. This was followed by unclassified_g_Glomus_f_Glomeraceae (unclassified Glomus taxa belonging to the family Glomeraceae), which contributed 20.63% of the total AMF community. In contrast, Glomus sp. VTX00304 had the lowest relative abundance, making up only 0.05% of the total.
The Glomus wirsel OTU6 (VTX00202) exhibited the highest relative abundance, primarily distributed in the rhizosphere soils associated with Arundinella anomala, Artemisia mongolica, and Clematis hexapetala. Additionally, this population was also detected in soil samples associated with Artemisia anethifolia and present at lower proportions in the rhizosphere soils of Taraxacum mongolicum and Puccinellia tenuiflora. In contrast, Glomus MO G3 (VTX00113) was predominantly found in the rhizosphere soils of Leymus chinensis and Vicia amoena, whereas Glomus lamellosum (VTX00193) was more commonly observed in the rhizosphere soils of Taraxacum mongolicum and Puccinellia tenuiflora.
Further analysis revealed plant-specific AMF communities in the rhizosphere soils of different host species. For instance, Glomus Franke A1 (VTX00076) was unique to the rhizosphere soil of Arundinella anomala; Glomus Glo E (VTX00319) and Glomus MO G14 (VTX00083) were exclusively detected in the rhizosphere soil of Taraxacum mongolicum; Glomus Douhan9 (VTX00056) and Glomus Glo16 (VTX00120) were unique to the rhizosphere soil of Puccinellia tenuiflora; Glomus acnaGlo2 (VTX00155) was specific to the rhizosphere soil of Artemisia mongolica; Glomus Glo D (VTX00103) occurred exclusively in the rhizosphere soil of Artemisia anethifolia; Glomus sp. (VTX00165) was unique to the rhizosphere soil of Clematis hexapetala, whereas Glomus caledonium (VTX00065) was a specific component of the rhizosphere soil of Vicia amoena (Figure 2).
The sample distance matrix was calculated using the Bray–Curtis distance algorithm and subjected to hierarchical clustering analysis (Figure 3), clearly revealing the proximity relationships among samples. The results showed that the rhizosphere soils of Vicia amoena and Taraxacum mongolicum each formed distinct clusters, indicating that these two plant species exhibit unique AMF species compositions and differ significantly from other plants.

3.4. Analysis of Rhizosphere pH, EC, and AMF Diversity in Eight Different Plant Species

Eight plant species showed significant differences in rhizosphere soil pH and EC (p < 0.05). Taraxacum mongolicum had the highest pH, followed by Arundinella anomala, Leymus chinensis, Clematis hexapetala, and Puccinellia tenuiflora—all significantly higher than low-pH species (Artemisia mongolica, Artemisia anethifolia, Vicia amoena), with Vicia amoena having the lowest pH. For EC, Arundinella anomala had the highest value, followed by Leymus chinensis, Taraxacum mongolicum, Clematis hexapetala, and Puccinellia tenuiflora. Arundinella anomala clear salinization gradient existed: high-salinity group (A–D, G) had significantly higher EC than low-salinity group (E, F, H), with Vicia amoena having the lowest EC.
For AMF, one-way ANOVA revealed highly significant differences in both spore density and hyphal length density among plant species (p < 0.001). Taraxacum mongolicum exhibited significantly higher spore density and hyphal length density compared to all other species (p < 0.05). In contrast, Vicia amoena displayed the lowest spore density, while Clematis hexapetala showed the lowest hyphal length density. With respect to species richness, Puccinellia tenuiflora recorded the highest value (15), exceeding the overall mean by 38.1%. Notably, Taraxacum mongolicum and Puccinellia tenuiflora exceeded the overall means across all three AMF parameters (Table 4).
Eight plant species exhibited significant differences in rhizosphere soil AMF community diversity indices (Chao 1, Shannon, Simpson, Sobs). Arundinella anomala had the highest Sobs and Chao 1 indices, followed by Taraxacum mongolicum and Puccinellia tenuiflora (both for Sobs/Chao 1), while Clematis hexapetala showed the lowest Sobs and Chao 1. For Shannon, Taraxacum mongolicum exhibited the highest value, followed by Puccinellia tenuiflora and Leymus chinensis (both labeled), with Clematis hexapetala being the lowest. Simpson (dominance) was highest in Clematis hexapetala and Arundinella anomala, and lowest in Taraxacum mongolicum and Puccinellia tenuiflora. Notably, the high EC group (A–D, G) had significantly higher mean Sobs (32.0 vs. 8.2) and Shannon (1.8 vs. 1.0) than the low EC group (E, F, H) (Figure 4a–d).

4. Discussion

4.1. Species and Distribution of Arbuscular Mycorrhizal Fungi (AMF) in the Rhizosphere of Eight Salt-Tolerant Plants in the Songnen Grassland

The Songnen Grassland, a representative saline–alkali ecosystem in Northeast China, is increasingly threatened by soil salinization, making the exploration of plant–AMF symbiotic mechanisms critical for ecological restoration. Here, we integrated morphological identification with high-throughput sequencing to characterize AMF diversity in rhizospheres of eight salt-tolerant plant species, uncovering the diversity, community structure, and environmental adaptability of AMF in their rhizospheres within the Songnen Grassland. This work offers critical theoretical support for applying salt-tolerant plant–AMF symbiotic systems to saline–alkali land restoration.
Morphological identification revealed 22 AMF species belonging to seven genera, with Glomus as the dominant genus (eight species, accounting for 36.4% of the total), consistent with previous studies on saline–alkali grasslands [39,40]. High-throughput sequencing further confirmed the dominance of Glomus, detecting 34 virtual taxa (VTs) that contributed to 85% of total sequences and 89.33% of relative abundance. This dominance is likely attributed to Glomus’s unique adaptations to salt stress [41], such as the formation of thick-walled spores for enhanced survival and efficient nutrient acquisition—traits that enable it to maintain high survival rates and reproductive efficiency in saline–alkali environments [42,43]. Among the identified species, Glomus mosseae was particularly prominent. Widespread species such as Glomus mosseae and Glomus etunicatum were detected across all samples, indicating their broad host affinity [44], while rare species such as Glomus deserticola (only found in the rhizosphere of Taraxacum mongolicum) suggested potential host specificity or niche specialization [45].
Discrepancies between morphological and molecular approaches underscore the complementary nature of these methodologies [46]: morphological identification proved highly effective in resolving species-level diversity (e.g., Glomus etunicatum), whereas high-throughput sequencing enabled the detection of non-sporulating or low-abundance taxa (e.g., Paraglomus brasilianum). For instance, Acaulospora was identified through morphological analysis but not recovered via sequencing, a discrepancy potentially attributable to primer bias (e.g., underamplification associated with NS31/AM1 primers) or suboptimal DNA extraction efficiency. These findings highlight the necessity of integrating multiple methodological approaches to achieve a comprehensive characterization of AMF communities [46].
Beyond the Songnen Grassland, Glomus has been frequently reported as a dominant genus in diverse ecosystems (e.g., tropical forests, temperate grasslands, agricultural systems), a phenomenon linked to its exceptional adaptability [47]. Its thick-walled spores enhance tolerance to adverse conditions, while optimal concentrations of soil organic matter, nitrogen, and phosphorus further support its growth [48]. Ecologically, Glomus exhibits higher efficiency in establishing and maintaining symbiotic relationships with plant roots compared to other AMF genera [49]. Its high abundance and diversity improve plant utilization of limited resources—particularly under low-nutrient or extreme conditions—and enhance stress resistance. Additionally, Glomus regulates soil microbial community structure by secreting extracellular enzymes that decompose complex organic compounds (releasing plant-available mineral nutrients) and forming synergies with other microorganisms, thereby optimizing the soil microecosystem [50].
Glomus mosseae and Glomus etunicatum were isolated from all soil samples, indicating their broad distribution and strong affinity for the selected plant species [41]. Other species (e.g., Glomus constrictum, Acaulospora laevis, Rhizophagus intraradices) were present in five samples, representing common AMF types in the Songnen Grassland. In contrast, Glomus deserticola was only detected in a single sample (rhizosphere of Puccinellia tenuiflora), making it a rare type. These patterns reflect significant differences in AMF communities between plant rhizospheres or across environments—a phenomenon widely reported in global ecosystems [51]. Through comparison with the MaarjAM database, Glomus versiforme OTU6 VTX00202 was found to have the highest relative abundance in our study and was also distributed in the rhizosphere of Magnoliaceae, Liliaceae, and Poaceae plants, highlighting its broad ecological niche [52,53]. Both morphological and molecular methods consistently identified Glomus as the dominant genus in the rhizosphere of multiple plant species. This consistency supports the conclusion that Glomus’s wide distribution across ecosystems is driven by its strong environmental adaptability and metabolic flexibility, allowing it to survive and function under diverse soil types and climatic conditions worldwide [54].
Although Glomus dominated rhizosphere AMF communities in the Songnen saline–alkali grassland due to high abundance and wide distribution, the ecological significance of low-abundance genera (e.g., Acaulospora, Rhizophagus, Ambispora, Archaeospora, Gigaspora, Diversispora) should not be overlooked [40,55]—they play critical roles in niche differentiation, functional complementarity, and ecosystem stability under extreme conditions (e.g., high HCO3, low nutrients) [56,57]. Acaulospora (in Leymus chinensis/Puccinellia tenuiflora rhizospheres) uses thick spores/branched hyphae to exploit insoluble organic phosphorus (e.g., phytate) via phytase, complementing Glomus’s inorganic phosphorus reliance [52,58]; Rhizophagus (R. intraradices in Taraxacum mongolicum/Vicia amoena) forms tight cortical symbiosis to regulate ion balance (increased K+, reduced Na+), enhancing salt tolerance [59]; Ambispora (A. leptoticha in Artemisia mongolica) secretes polysaccharides to promote microaggregates (<0.25 mm), ameliorating compaction [60]. Through niche differentiation (organic vs. inorganic P, surface vs. deep soil) and functional complementarity (ion regulation, soil improvement), these genera enhance AMF functional diversity and ecosystem resilience [61].

4.2. Adaptive Strategies of AMF–Plant Interactions in Rhizospheres of Different Plants in Heterogeneous Habitats

AMF exhibit weak host specificity and facilitate resource redistribution (e.g., carbon, nitrogen, phosphorus) among plants via extensive extraradical hyphal networks [12], thereby modulating interspecific relationships and community composition [62]. This resource-exchange mechanism promotes nutrient sharing among plant individuals, alleviates stress responses to adverse conditions (e.g., drought or nutrient deficiency), and ultimately contributes to the structural and functional stability of ecosystems [34]. Additionally, AMF play a key role in soil aggregate formation, enhancing soil organic matter content and carbon sequestration—processes critical to the global carbon cycle [63]. Deepening our understanding of the interactions between AMF, plants, and their environment holds profound value for sustainable agriculture and ecosystem management [15].
Plant–soil microbial interactions serve as a critical link between aboveground and belowground components of terrestrial ecosystems, where changes in aboveground plant diversity influence the structure and function of soil microbial communities [62]. Although AMF are widely distributed, their communities are often considered to have low species diversity and weak host specificity [64]. However, advances in molecular techniques and growing recognition of AMF functional diversity have revealed that their species diversity and genetic variation may be severely underestimated, and their host specificity requires re-evaluation. AMF diversity is more closely linked to plant diversity than to soil bacterial communities [55], and studies have shown that grassland plants form specific associations with their rhizosphere AMF communities—even as plant–AMF relationships are generally regarded as non-specific [35].
Our study confirmed specific relationships between different grassland plant species and AMF, with significant differences in soil properties among the rhizospheres of the eight studied plants. This indicates that each plant species contributes uniquely to belowground ecosystem function. Rhizosphere AMF diversity indices not only reflect host–AMF affinity but also reveal the complex ecological relationships shaped by long-term coevolution. AMF spore distribution in rhizosphere soils showed marked spatial heterogeneity, with significant variation in spore density, hyphal density, and abundance across plots. These patterns are jointly driven by host plant growth cycles, soil physicochemical properties, moisture conditions, and AMF reproductive strategies, leading to distinct temporal and spatial heterogeneity in AMF communities [65].
Dynamic changes in the rhizosphere microenvironment further shape AMF community structure: fluctuations in soil pH regulate interspecific competition among AMF, while changes in organic matter content alter their carbon source acquisition [35]. Plant exudates—varying in composition and spatiotemporal distribution—further modulate AMF community assembly [27]. This multi-level interaction reflects the tight coupling of rhizosphere biotic and abiotic factors and underscores the complexity of AMF dynamics [66]. Soil microenvironment and plant species differences determine the specificity of plant–AMF relationships in grasslands, and future research should explore complementary or competitive interactions among microbial populations [67].
Our study found significant differences in rhizosphere soil pH and EC among plants: Taraxacum mongolicum had the highest pH, Arundinella anomala had the highest EC, and Vicia amoena had the lowest EC. Plots formed clear gradients: groups A–D, G (high salinization) and groups E, F, H (low salinization). Previous studies identify EC and pH as major drivers of AMF community composition in Songnen saline–alkali grassland vegetation succession [39], and our results confirm that salt-tolerant plants here form multi-dimensional saline–alkali adaptation mechanisms via synergistic differentiation of rhizosphere microenvironment (pH, EC) and AMF symbiotic strategies. The significant differences in AMF community diversity indices primarily stemmed from two key interacting drivers: First, plant species-specific interactions [68]. Different plant species produce distinct root exudates that selectively modulate AMF colonization. For instance, Taraxacum mongolicum and Puccinellia tenuiflora release compounds like citric acid and sucrose, which enhance AMF spore germination and hyphal growth [45]. In contrast, Clematis hexapetala and Vicia amoena secrete phenolic compounds that suppress AMF establishment [58]. These species-specific exudate profiles directly shape the composition and diversity of AMF communities in the rhizosphere [69]. Second, soil physicochemical gradients. High-salinity (EC: 2.07–2.61 mS/cm) and high-pH (8.70–8.81) conditions favored the survival of salt- and alkali-tolerant AMF taxa (e.g., Glomus, Claroideoglomus), which are adapted to acquiring nutrients under stressful conditions. Conversely, acidic (pH: 7.30–7.49) and low-salinity (EC: 0.20–0.48 mS/cm) soils limited the diversity of most AMF taxa, as many species lack tolerance to acidic environments [39]. These two factors acted synergistically to drive the pronounced variation in arbuscular mycorrhizal fungal community diversity observed across plant species.
During symbiosis establishment, AMF’s selective preference for host plants is a key driver of community structure and distribution—shaped by host genetic background, root morphology, metabolic activity, and environmental conditions [70]. This preference directly regulates critical AMF physiological processes (e.g., spore germination, hyphal expansion, colonization) and indirectly influences plant growth [3], biomass allocation, and stress resistance by modulating nutrient (phosphorus, nitrogen) uptake [62]. The synergy between AMF preference and root nutrient demands is precisely regulated by root exudate types and concentrations, while root-released signal molecules induce spore germination and hyphal chemotaxis toward roots [66]. Besides the key driving roles of pH and EC in AMF community structure examined in this study, environmental factors such as soil moisture, organic matter content, and temperature may also significantly affect AMF diversity and distribution [34,71,72]. Soil moisture directly regulates the microenvironment for AMF spore germination and hyphal extension, with high moisture facilitating continuous hyphal network expansion and drought potentially limiting spore activity [73]. The content differences of organic matter, as a supplementary carbon source besides host photosynthates, alter the resource competition pattern among AMF species, with organic matter-preferring species more likely to become dominant in high-content soils [74]. Temperature affects AMF metabolic activity, with optimal temperatures enhancing growth and reproduction rates and extreme temperatures potentially inhibiting their functions [71]. Meanwhile, AMF play important roles in saline–alkali grassland ecosystems: the dense hyphal network of the dominant genus Glomus can expand the host’s absorption range of nutrients such as phosphorus and nitrogen [41], promoting effective nutrient cycling in saline–alkali soils [75]; specialized species like G. deserticola, only detected in the rhizosphere of Taraxacum mongolicum, may enhance plant resistance to salt stress by regulating host root ion balance or secreting stress-resistant metabolites [45]; high-diversity AMF communities maintain ecosystem stability through functional complementarity, with different species participating in processes such as nutrient cycling and soil structure improvement [76].
Understanding these mechanisms is essential for elucidating terrestrial ecosystem stability and function. Most AMF in the Songnen grassland exhibit strong adaptability to saline–alkaline soils, with a pronounced rhizosphere distribution, underscoring their indispensable role in maintaining the structure and function of saline–alkaline ecosystems. Systematic screening and validation of suitable AMF combinations, coupled with targeted soil amelioration, can effectively enhance host plant growth, optimize the soil microenvironment, and improve ecological restoration efficiency in the Songnen saline–alkaline grassland [36]. AMF facilitate nutrient uptake and enhance stress tolerance through efficient symbiotic associations, providing critical insights into microbial–plant symbiotic relationships in complex ecosystems.
Taraxacum mongolicum exhibited the highest AMF diversity, likely due to root exudates that attract diverse, salt-tolerant Glomus species—thereby enhancing mitigation of ion toxicity and improving nutrient uptake [72]. In contrast, plants from low-salt habitats, such as Clematis hexapetala (lowest Sobs index), exhibited simplified AMF communities, relying on efficient Glomus species to optimize resource allocation and minimize symbiotic costs, thus maintaining stable symbiosis under nutrient-limited conditions [75]. High-salinity plants (Arundinella anomala, Taraxacum mongolicum) enhance AMF symbiosis to balance carbon consumption and salt stress, whereas low-salinity plants (Artemisia mongolica, Vicia amoena) shift toward functional degeneracy or non-symbiotic resistance [77]. This bidirectional plant–AMF regulation reveals dynamic coevolution in heterogeneous habitats, providing a niche-based theoretical foundation and practical support for screening salt-tolerant AMF and optimizing mycorrhizal technology.
Hierarchical clustering and OTU distribution analysis revealed distinct AMF communities associated with Taraxacum mongolicum and Vicia amoena, confirming plant species as a key driver of AMF community structure—consistent with global patterns in grasslands and forests [78]. Unique VTs (e.g., Glomus Franke A1 in Arundinella anomala, Glomus caledonium in Vicia amoena) further support host specificity, likely shaped by root exudates, nutrient demands, and evolutionary history [66]. A critical finding was that AMF diversity in the high EC group (A–D, G) was significantly higher than in the low EC group (E–F, H), with mean Sobs and Shannon indices clearly differentiating the two groups. This contradicts the common assumption that high salinity reduces microbial diversity, instead indicating that AMF in salt-tolerant plant rhizospheres have adapted to—even thrive in—high-salt environments [79]. The positive EC-AMF diversity correlation may reflect a “stress-induced symbiosis” mechanism, where saline–soil plants rely more heavily on AMF for nutrient acquisition (e.g., phosphorus) and salt tolerance.

5. Conclusions

Through a systematic analysis of AMF in the rhizosphere of eight saline–alkali-tolerant plant species in the Songnen Grassland, this study confirmed that AMF communities exhibit unique ecological adaptation mechanisms in high saline–alkali habitats. Morphological identification revealed 22 AMF species (belonging to 8 genera), with Glomus (36.4%) and Acaulospora (22.7%) as the dominant genera; high-throughput sequencing detected 40 virtual taxa (VTs), where Glomus accounted for an overwhelming 89.33% relative abundance. This discrepancy stems from the combined effects of primer bias and detection limits for low-abundance species, underscoring the necessity of using multiple methods in conjunction. Rhizosphere soil EC showed a significant gradient. The Shannon index of AMF in the high-salt group was higher than that in the control group, indicating that salinization did not inhibit AMF diversity but instead activated their salt-tolerant physiological response networks. Hierarchical clustering analysis showed that the AMF community structure in the rhizosphere of Taraxacum mongolicum and Vicia amoena deviated significantly from other species. This differentiation arises from differences in the interaction between plant root exudates and mycorrhizal colonization strategies, confirming that host species are the core biological factor in AMF community assembly. These findings provide a theoretical basis for the ecological restoration of saline–alkali lands: selecting Glomus strains and constructing symbionts with specific hosts can enhance the efficiency of vegetation reconstruction by strengthening mycorrhizal networks. Our study elucidates the dynamic coevolutionary relationships between salt-tolerant plants and AMF in the Songnen saline–alkali grassland, highlighting AMF’s critical role in enhancing plant adaptation to saline–alkali stress. These findings provide valuable insights into the ecological mechanisms underlying plant–AMF symbiosis in heterogeneous habitats and offer practical guidance for selecting salt-tolerant AMF strains and optimizing ecological restoration strategies for saline–alkali ecosystems.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/agronomy15092070/s1, Figure S1: Geographic location of the study site and distribution of rhizosphere soil sampling points in the Songnen Grassland, Northeast China; Figure S2: Optical micrographs of arbuscular mycorrhizal fungal spores identified through morphological characterization.

Author Contributions

Conceptualization, L.M.; methodology, Y.L. and T.J.; software, Z.W.; validation, X.Y.; formal analysis, T.J.; investigation, L.M.; resources, Z.W. and Y.W.; data curation, L.M.; writing—original draft preparation, Y.W. and X.Y.; writing—review and editing, L.M. and Y.L.; visualization, Y.W.; supervision, Z.W., Z.X. and T.J.; project administration, L.M. and X.Y.; funding acquisition, L.M. and X.Y. All authors have read and agreed to the published version of the manuscript.

Funding

This research was supported by the National Natural Science Foundation of China (32201469, 32101396); X.Y. acknowledges support by the Innovation Leading Talents Project of the “Tianchi Talent” Recruitment Plan of Xinjiang Uygur Autonomous Region.

Data Availability Statement

The data supporting the findings of this study are available from the corresponding author upon reasonable request.

Acknowledgments

We would like to thank the anonymous reviewers for helpful comments on the manuscript.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. AMF species composition in rhizospheres of different plant species at the species level.
Figure 1. AMF species composition in rhizospheres of different plant species at the species level.
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Figure 2. Bidirectional semicircular plot showing associations between AMF community composition in rhizospheres of eight salt-tolerant plants and their host distribution. Left semicircle (plants): Each sector represents a plant rhizosphere sample (A–H, Table 2). Colored segments indicate relative abundances of AMF virtual taxa (VTs; MaarjAM database, 97% sequence similarity). Right semicircle (AMF): Each sector represents an AMF VT. Colored segments indicate distribution of the VT across plant rhizospheres (colors match left semicircle).
Figure 2. Bidirectional semicircular plot showing associations between AMF community composition in rhizospheres of eight salt-tolerant plants and their host distribution. Left semicircle (plants): Each sector represents a plant rhizosphere sample (A–H, Table 2). Colored segments indicate relative abundances of AMF virtual taxa (VTs; MaarjAM database, 97% sequence similarity). Right semicircle (AMF): Each sector represents an AMF VT. Colored segments indicate distribution of the VT across plant rhizospheres (colors match left semicircle).
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Figure 3. Hierarchical clustering of AMF community structure in rhizospheres of eight salt-tolerant plants. UPGMA clustering based on Bray–Curtis dissimilarity.
Figure 3. Hierarchical clustering of AMF community structure in rhizospheres of eight salt-tolerant plants. UPGMA clustering based on Bray–Curtis dissimilarity.
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Figure 4. AMF diversity indices in rhizospheres of eight salt-tolerant plants: Chao1 index (a); Shannon index (b); Simpson index (c); Sobs index (d). Different lowercase letters indicate significant differences among plant species (p < 0.05).
Figure 4. AMF diversity indices in rhizospheres of eight salt-tolerant plants: Chao1 index (a); Shannon index (b); Simpson index (c); Sobs index (d). Different lowercase letters indicate significant differences among plant species (p < 0.05).
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Table 1. Taxonomic composition and species diversity of AMF in rhizospheres of plants.
Table 1. Taxonomic composition and species diversity of AMF in rhizospheres of plants.
GenusSpecies
GlomusG. deserticola, G. mosseae, G. pansihalos, G. convolutum, G. magnicaule, G. etunicatum, G. tenebrosum, Glomus sp. 1
AcaulosporaA. foveata, A. laevis, A. lacunosa, A. mellea, A.rehmii
RhizophagusR. Fasciculatus, R. manihotis, R. intraradices, R. cluram
AmbisporaA. Leptoticha, A. gerdemannii
ClaroideoglomusC. claroideum
GigasporaGi. decipiens
FunneliformisF. constrictum
Table 2. Virtual taxa of inter-rhizosphere arbuscular mycorrhizal fungi associated with different plant species identified through high-throughput sequencing. The entries in the table represent the genus names and corresponding virtual species numbers of arbuscular mycorrhizal fungi detected in the rhizosphere of various plants.
Table 2. Virtual taxa of inter-rhizosphere arbuscular mycorrhizal fungi associated with different plant species identified through high-throughput sequencing. The entries in the table represent the genus names and corresponding virtual species numbers of arbuscular mycorrhizal fungi detected in the rhizosphere of various plants.
GenusVirtual Taxa
GlomusGlomus Franke A1 VTX00076, Glomus Franke A1 VTX00269, Glomus Glo C VTX00323, Glomus Glo D VTX00103, Glomus Glo E VTX00319, Glomus Glo7 VTX00214, Glomus Glo16 VTX00120, Glomus PSAMG1 VTX00291, Glomus Yamato2005 D VTX00084, Glomus Yamato2005 D VTX00224, Glomus sp. VTX00165
Glomus sp. VTX00279, Glomus sp. VTX00304, Glomus caledonium VTX00065, Glomus Douhan9 VTX00056, Glomus lamellosu VTX00193, Glomus perpusillum VTX00287
Glomus GlAc3.1 VTX00190, Glomus GlAd3.3 VTX00289, Glomus MO G3 VTX00113
Glomus MO G4 VTX00166, Glomus MO G13 VTX00115, Glomus MO G14 VTX00083
Glomus MO G16 VTX00072, Glomus MO G22 VTX00125, Glomus Wirsel OTU12 VTX00188, Glomus Wirsel OTU6 VTX00202, Glomus acnaGlo2 VTX00155
Glomus acnaGlo7 VTX00057, Glomus viscosum VTX00063, Glomus ORVIN GLO3B VTX00223, Glomus ORVIN GLO3E VTX00309, Glomus ORVIN GLO3D VTX00310
Glomus ORVIN GLO4 VTX00278
ArchaeosporaArchaeospora Other1 VTX00005, Archaeospora sp. VTX00009
ParaglomusParaglomus brasilianum VTX00239, Paraglomus occultum VTX00238
DiversisporaDiversispora MO GC1 VTX00060
GigasporaGigaspora decipiens VTX00039
Acaulospora ------
Table 3. Distribution of AMF in the rhizosphere of different plants. A: Arundinella anomala; B: Leymus chinensis; C: Taraxacum mongolicum; D: Puccinellia tenuiflora; E: Artemisia mongolica; F: Artemisia anethifolia; G: Clematis hexapetala; H: Vicia amoena.
Table 3. Distribution of AMF in the rhizosphere of different plants. A: Arundinella anomala; B: Leymus chinensis; C: Taraxacum mongolicum; D: Puccinellia tenuiflora; E: Artemisia mongolica; F: Artemisia anethifolia; G: Clematis hexapetala; H: Vicia amoena.
GenusSpecieABCDEFGH
GlomusG. deserticola
G. mosseae
G. pansihalos
G. convolutum
G. magnicaule
G. etunicatum
G. tenebrosum
Glomus sp. 1
AcaulosporaA. foveata
A. laevis
A. lacunosa
A. mellea
A.rehmii
RhizophagusR. fasciculatus
R. manihotis
R. intraradices
R. cluram
AmbisporaA. leptoticha
A.gerdemannii
ClaroideoglomusC. claroideum
GigasporaGi. decipiens
FunneliformisF. constrictum
Table 4. Rhizosphere AM fungal abundance indices of different plant species.
Table 4. Rhizosphere AM fungal abundance indices of different plant species.
PlantspHElectrical Conductivity (mS/cm)Spore Density
(Number per Gram)
Hyphal Length Density (m/g Dry Soil)Species
Richness
A8.79 ± 0.04 ab2.61 ± 0.06 a12.0 ± 0.69 ab6.38 ± 0.37 d10
B8.76 ± 0.04 ab2.49 ± 0.03 ab10.67 ± 0.62 ab7.22 ± 0.42 c12
C8.81 ± 0.07 a2.43 ± 0.05 b15.0 ± 0.87 a16.94 ± 0.98 a13
D8.70 ± 0.06 b2.07 ± 0.04 d13.33 ± 0.77 ab13.01 ± 0.75 b15
E7.49 ± 0.05 c0.48 ± 0.01 e8.67 ± 0.50 b6.67 ± 0.39 d9
F7.35 ± 0.04 d0.40 ± 0.02 e7.67 ± 0.44 b8.88 ± 0.51 c11
G8.77 ± 0.06 ab2.30 ± 0.18 c4.67 ± 0.27 c3.94 ± 0.23 e7
H7.30 ± 0.07 e0.20 ± 0.01 f4.0 ± 0.23 c5.88 ± 0.34 d10
Note: Table values represent mean ± standard deviation. Different lowercase letters denote significant differences in AMF abundance among different plant soil samples (p < 0.05)
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Mei, L.; Liu, Y.; Wang, Z.; Xiong, Z.; Wang, Y.; Jin, T.; Yang, X. Diversity and Community Structure of Rhizosphere Arbuscular Mycorrhizal Fungi in Songnen Grassland Saline–Alkali-Tolerant Plants: Roles of Environmental Salinity and Plant Species Identity. Agronomy 2025, 15, 2070. https://doi.org/10.3390/agronomy15092070

AMA Style

Mei L, Liu Y, Wang Z, Xiong Z, Wang Y, Jin T, Yang X. Diversity and Community Structure of Rhizosphere Arbuscular Mycorrhizal Fungi in Songnen Grassland Saline–Alkali-Tolerant Plants: Roles of Environmental Salinity and Plant Species Identity. Agronomy. 2025; 15(9):2070. https://doi.org/10.3390/agronomy15092070

Chicago/Turabian Style

Mei, Linlin, Yingbin Liu, Zixian Wang, Zixuan Xiong, Yuze Wang, Tianqi Jin, and Xuechen Yang. 2025. "Diversity and Community Structure of Rhizosphere Arbuscular Mycorrhizal Fungi in Songnen Grassland Saline–Alkali-Tolerant Plants: Roles of Environmental Salinity and Plant Species Identity" Agronomy 15, no. 9: 2070. https://doi.org/10.3390/agronomy15092070

APA Style

Mei, L., Liu, Y., Wang, Z., Xiong, Z., Wang, Y., Jin, T., & Yang, X. (2025). Diversity and Community Structure of Rhizosphere Arbuscular Mycorrhizal Fungi in Songnen Grassland Saline–Alkali-Tolerant Plants: Roles of Environmental Salinity and Plant Species Identity. Agronomy, 15(9), 2070. https://doi.org/10.3390/agronomy15092070

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