放牧干扰下丛枝菌根真菌群落对土壤多功能性的影响

doi:10.17521/cjpe.2025.0117

植物生态学报 ›› 2026, Vol. 50 ›› Issue (3): 685-699.DOI: 10.17521/cjpe.2025.0117 cstr: 32100.14.cjpe.2025.0117

放牧干扰下丛枝菌根真菌群落对土壤多功能性的影响

江康威¹, 吕程², 王亚菲³, 李宏¹, 张芷晴², 王雨², 张青青²^,^*(), 吐尔逊娜依•热依木²

¹ 新疆农业大学草业学院, 乌鲁木齐 830052
² 新疆农业大学生命科学学院新疆极端环境生物生态适应与进化重点实验室, 乌鲁木齐 830052
³ 新疆农业大学资源与环境学院, 乌鲁木齐 830052

收稿日期:2025-04-01 接受日期:2025-07-10 出版日期:2026-03-20 发布日期:2026-04-07
通讯作者: *张青青(greener2010@sina.com)
基金资助:
国家自然科学基金(32260311);国家自然科学基金(41561103)

Effects of soil arbuscular mycorrhizal fungal communities on soil multifunctionality under grazing disturbance

JIANG Kang-Wei¹, LÜ Cheng², WANG Ya-Fei³, LI Hong¹, ZHANG Zhi-Qing², WANG Yu², ZHANG Qing-Qing²^,^*(), Tuerxunnayi REYIMU²

¹ College of Grassland Science, Xinjiang Agricultural University, Ürümqi 830052, China
² Xinjiang Key Laboratory for Ecological Adaptation and Evolution of Extreme Environment Biology, College of Life Sciences, Xinjiang Agricultural University, Ürümqi 830052, China
³ College of Resources and Environment, Xinjiang Agricultural University, Ürümqi 830052, China

Received:2025-04-01 Accepted:2025-07-10 Online:2026-03-20 Published:2026-04-07
Contact: *ZHANG Qing-Qing(greener2010@sina.com)
Supported by:
National Natural Science Foundation of China(32260311);National Natural Science Foundation of China(41561103)

摘要/Abstract

摘要：

丛枝菌根真菌(AMF)与植物根系的共生合作关系改善了植物群落的养分吸收效率, 对维持草地生态系统关键的功能和过程起到至关重要的作用。然而, 不同放牧强度下AMF群落的变化规律仍存在争议, 特别是在放牧干扰下AMF群落对土壤多功能性(SMF)的驱动作用仍未被明确解析。该研究选取新疆天山北坡的未放牧、轻度放牧和重度放牧的山地草甸草原为研究对象, 采用高通量测序技术分析AMF群落并构建共现网络。采用基于聚类分析的加权平均值法系统评估SMF, 探究不同放牧强度下AMF群落特征及其对SMF的驱动作用。结果表明, 3个放牧强度的AMF优势属均为球囊菌属(Glomus)和多样孢囊霉属(Diversispora)。随着放牧强度增加, AMF群落的多样性和共现网络的稳定性、复杂性、内聚力以及SMF均呈现先增加后降低的单峰曲线趋势。线性回归和结构方程模型的结果表明, 放牧可通过调控AMF群落多样性、网络复杂性和稳定性驱动SMF。AMF群落多样性对SMF的驱动效应主要依靠其网络复杂性和稳定性来实现, 且群落多样性的直接效应低于复杂性和稳定性。该研究强调AMF群落网络复杂性和稳定性放大了群落多样性对SMF的驱动效应, 强化了群落多样性与SMF的联系。研究结果为实现退化草地生态系统的恢复和可持续管理提供理论参考。

关键词: 丛枝菌根真菌, 共现网络分析, 稳定性, 网络复杂性, 土壤多功能性, 放牧, 山地草甸草原

Abstract:

Aims The symbiotic relationship between arbuscular mycorrhizal fungi (AMF) and plant roots enhances nutrient acquisition in plant communities and plays a vital role in sustaining key functions and ecological processes in grassland ecosystems. However, the response patterns of AMF communities to varying grazing intensities remain under debate, and the mechanisms by which these communities influence soil multifunctionality (SMF) under grazing disturbance are not yet fully understood.

Methods To address these knowledge gaps, this study investigated mountain meadow steppes on the northern slope of the Tianshan Mountains in Xinjiang under three grazing intensities: ungrazed, lightly grazed, and heavily grazed. High-throughput sequencing was used to characterize the composition of AMF communities and to construct their co-occurrence networks. A weighted average method, based on cluster analysis, was applied to comprehensively evaluate SMF and to analyze how AMF community characteristics vary with grazing intensities and how these variations influence SMF.

Important findings The results indicated that the dominant AMF genera in the three grazing treatments were Glomus and Diversispora. As grazing intensity increased, AMF community diversity, as along with network stability, complexity, cohesion, and SMF, exhibited a unimodal pattern—increasing initially and then declining at higher grazing levels. Linear regression and structural equation modeling indicated that grazing influences SMF mainly by modulating AMF community diversity and altering the complexity and stability of AMF networks. The effect of AMF community diversity on SMF was largely indirect, mediated through its effects on network structure, while its direct impact was relatively weak. Overall, this study highlights that the complexity and stability of AMF co-occurrence networks enhance the role of community diversity in shaping SMF and strengthen the interdependent relationship between biodiversity and ecosystem functioning. These findings offer valuable theoretical insights for the restoration and sustainable management of degraded grassland ecosystems.

Key words: arbuscular mycorrhizal fungi, co-occurrence network analysis, stability, network complexity, soil multifunctionality, grazing, mountain meadow steppes

江康威, 吕程, 王亚菲, 李宏, 张芷晴, 王雨, 张青青, 吐尔逊娜依•热依木. 放牧干扰下丛枝菌根真菌群落对土壤多功能性的影响. 植物生态学报, 2026, 50(3): 685-699. DOI: 10.17521/cjpe.2025.0117

JIANG Kang-Wei, LÜ Cheng, WANG Ya-Fei, LI Hong, ZHANG Zhi-Qing, WANG Yu, ZHANG Qing-Qing, Tuerxunnayi REYIMU. Effects of soil arbuscular mycorrhizal fungal communities on soil multifunctionality under grazing disturbance. Chinese Journal of Plant Ecology, 2026, 50(3): 685-699. DOI: 10.17521/cjpe.2025.0117

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2025.0117

https://www.plant-ecology.com/CN/Y2026/V50/I3/685

图/表 6

表1 天山北坡中段山地草甸不同放牧样地中的植物群落组成(平均值±标准误)

Table 1 Plant community composition in different grazing treatments in the mountain meadow steppes on the northern slope at middle section of the Tianshan Mountains (mean ± SE)

放牧强度 Grazing intensity	植物优势种 Dominant plant species	高度 Height (cm)	盖度 Coverage (%)	地上生物量 Aboveground biomass (g·m^-2)
对照 No grazing	针茅、羊茅、草地早熟禾 Carex stenocarpa, Festuca ovina, Poa pratensis	26.18 ± 3.63	87.50 ± 2.21	265.29 ± 60.24
轻度放牧 Light grazing	细果薹草、针茅、无芒雀麦 Carex stenocarpa, Stipa capillata, Bromus inermis	22.68 ± 1.77	93.50 ± 2.73	175.35 ± 19.29
重度放牧 Heavy grazing	醉马草、细果薹草、平车前 Achnatherum inebrians, Carex stenocarpa, Plantago depressa	13.21 ± 2.49	59.16 ± 3.67	123.31 ± 33.21

图1 天山北坡中段山地草甸未放牧、轻度放牧和重度放牧样地的位置。

Fig. 1 Location of ungrazed, lightly grazed and heavily grazed plots in the mountain meadow steppes on the northern slope at middle section of the Tianshan Mountains. CK, no grazing; HG, heavy grazing; LG, light grazing.

图2 天山北坡中段山地草甸不同放牧样地中运算分类单元(OTU)稀释曲线(A)、覆盖度指数(B)和数目(C) (平均值±标准误)的变化。CK, 未放牧; HG, 重度放牧; LG, 轻度放牧。不同小写字母代表不同放牧样地间差异显著(p < 0.05)。

Fig. 2 Changes in operational taxonomic unit (OTU) rarefaction curve (A), coverage index (B) and number (C) (mean ± SE) in different grazing treatments in the mountain meadow steppes on the northern slope at middle section of the Tianshan Mountains. CK, no grazing; HG, heavy grazing; LG, light grazing. Different lowercase letters indicate significant differences among different grazing treatments (p < 0.05).

图3 不同放牧样地中丛枝菌根真菌群落α多样性(A)、β多样性(B)、组成(C)和结构(D)的变化。CK, 未放牧; HG, 重度放牧; LG, 轻度放牧。Similarity, 相似性(1 - Total β); Total β, 总β多样性; Turnover β, 周转β多样性(物种替代过程); Nestedness β, 嵌套β多样性(丰富度差异过程)。ADONIS, 置换多元方差分析; ANOSIM, 相似性分析; NMDS1/NMDS2, 非度量多维尺度分析轴1/轴2; Stress, 胁迫系数。不同小写字母代表不同放牧样地间差异显著(p < 0.05)。

Fig. 3 Changes in alpha diversity (A), beta diversity (B), composition (C), and structure (D) of arbuscular mycorrhizal fungal communities in different grazing treatments. CK, no grazing; HG, heavy grazing; LG, light grazing. Similarity, 1 - Total β; Total β, total β diversity; Turnover β, turnover β diversity (replacement); Nestedness β, nestedness β diversity (richness difference). ADONIS, permutational multivariate analysis of variance; ANOSIM, similarity analysis; NMDS1/NMDS2, non-metric multidimensional scaling analysis axis 1/axis 2; Stress, stress coefficient. Different lowercase letters indicate significant differences among different grazing treatments (p < 0.05).

图4 不同放牧样地中丛枝菌根真菌群落共现网络特征的变化。A, 共现网络特征。B, 基于共现网络的关键种。图中虚线表示Zi ≥ 2.5或Pi ≥ 0.62的节点为关键种。C, 共现网络拓扑属性。D, 正内聚力。E, 负内聚力。F, 共现网络复杂性。G, 稳定性。H, 网络复杂性与稳定性的关系。CK, 未放牧; HG, 重度放牧; LG, 轻度放牧。不同小写字母代表不同放牧样地间差异显著(p < 0.05)。

Fig. 4 Changes in the co-occurrence network of arbuscular mycorrhizal fungal communities in different grazing treatments. A, Co-occurrence network characteristics. B, Key species based on co-occurrence network. Dashed lines indicate that nodes with Zi ≥ 2.5 or Pi ≥ 0.62 are identified as key species. C, Co-occurrence network topological properties. D, Positive cohesion. E, Negative cohesion. F, Co-occurrence network complexity. G, Stability. H, Relationship between network complexity and stability. CK, no grazing; HG, heavy grazing; LG, light grazing. Different lowercase letters indicate significant differences among different grazing treatments (p < 0.05).

图5 不同放牧样地中丛枝菌根真菌群落对土壤多功能性的驱动。A, 土壤多功能性。B, 基于随机森林的土壤多功能性的驱动因素。C, 土壤多功能性与丛枝菌根真菌群落多样性的关系。D, 土壤多功能性与丛枝菌根真菌群落共现网络复杂性和稳定性的关系。E, 基于随机森林土壤多功能性的影响因子。F, 基于结构方程模型的土壤多功能性驱动过程。G, 基于结构方程模型驱动因子对土壤多功能性的效应值。AB, 地上生物量; Ace index, Ace指数; AP, 碱性磷酸酶活性; Av-K, 速效钾含量; Av-N, 速效氮含量; Av-P, 速效磷含量; BB, 地下生物量; BD, 土壤密度; BG, β-1,4-葡萄糖苷酶活性; CBH, β-α-纤维二糖水解酶活性; Chao1 index, Chao1指数; Complexity, 网络复杂性; Coverage, 植物群落盖度; Density, 植物密度; EC, 电导率; Height, 植物群落高度; LAP, 亮氨酸氨基肽酶活性; MBC, 微生物生物量碳含量; MBN, 微生物生物量氮含量; MBP, 微生物生物量磷含量; NAG, β-1,4-N-乙酰氨基葡萄糖苷酶活性; Nestedness β, 嵌套β多样性; pH, 土壤pH; Shannon-Wiener index, Shannon-Wiener指数; Simpson index, Simpson指数; SOC, 土壤有机碳含量; Stability, 稳定性; SW, 土壤含水量; TK, 全钾含量; TN, 全氮含量; Total β, 总β多样性; TP, 全磷含量; Turnover β, 周转β多样性。CK, 未放牧; HG, 重度放牧; LG, 轻度放牧。不同小写字母代表不同放牧样地间差异显著(p < 0.05)。

Fig. 5 Arbuscular mycorrhizal fungal communities drive soil multifunctionality in different grazing treatments. A, Soil multifunctionality. B, Driving factors of soil multifunctionality based on random forest. C, Relationship between soil multifunctionality and the diversity of arbuscular mycorrhizal fungal communities. D, Relationship between soil multifunctionality and the complexity and stability of the co-occurrence network of arbuscular mycorrhizal fungal communities. E, Influencing factors based on the multifunctionality of soils in random forests. F, Driving process of soil multifunctionality based on structural equation model. G, Effect value of driving factors on soil multifunctionality based on structural equation model. AB, aboveground biomass; AP, alkaline phosphatase activity; Av-K, available potassium content; Av-N, available nitrogen content; Av-P, available phosphorus content; BB, belowground biomass; BD, soil density; BG, β-1,4-glucosidase activity; CBH, β-α-cellobiohydrolase activity; Coverage, coverage of plant community; Density, plant density; EC, electrical conductivity; Height, height of plant community; LAP, leucine aminopeptidase activity; MBC, microbial biomass carbon content; MBN, microbial biomass nitrogen content; MBP, microbial biomass phosphorus content; NAG, β-1,4-N-acetylglucosaminidase activity; Nestedness β, nested β diversity; pH, soil pH; SOC, soil organic carbon content; SW, soil water content; TK, total potassium content; TN, total nitrogen; Total β, total β diversity; TP, total phosphorus content; Turnover β, turnover β diversity. CK, no grazing; HG, heavy grazing; LG, light grazing. Different lowercase letters indicate significant differences among different grazing treatments (p < 0.05).

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放牧干扰下丛枝菌根真菌群落对土壤多功能性的影响

Effects of soil arbuscular mycorrhizal fungal communities on soil multifunctionality under grazing disturbance

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