丛枝菌根真菌对豆禾混播植物垂直生态位养分吸收的影响

doi:10.17521/cjpe.2025.0272

植物生态学报 ›› 2026, Vol. 50 ›› Issue (3): 760-773.DOI: 10.17521/cjpe.2025.0272 cstr: 32100.14.cjpe.2025.0272

丛枝菌根真菌对豆禾混播植物垂直生态位养分吸收的影响

魏莉¹, 王鹏森¹, 刘珊¹, 樊锐¹, 黄楠¹, 张建国¹, 其美拉姆², 苟扬¹, 刘沫含¹, 黄婷¹, 周冀琼¹^,^*()()

¹ 四川农业大学草业科技学院, 成都 611130
² 西藏拉萨市曲水县农业农村和水利局, 拉萨 850600

收稿日期:2025-07-21 接受日期:2026-02-25 出版日期:2026-03-20 发布日期:2026-05-20
通讯作者: *周冀琼(jiqiong_zhou@sicau.edu.cn)
基金资助:
国家自然科学基金(32271776);四川省自然科学基金(24NSFSC1813);四川省科技教育联合基金(25LHJJ0196);四川省科技教育联合基金(25LHJJ0190)

Arbuscular mycorrhizal fungi influence nutrient uptake along vertical niches in legume-grass mixtures

WEI Li¹, WANG Peng-Sen¹, LIU Shan¹, FAN Rui¹, HUANG Nan¹, ZHANG Jian-Guo¹, Qimeilamu ², GOU Yang¹, LIU Mo-Han¹, HUANG Ting¹, ZHOU Ji-Qiong¹^,^*()()

¹ College of Pratacultural Science, Sichuan Agricultural University, Chengdu 611130, China
² Qüxü County Bureau of Agriculture, Rural Affairs and Water Resources, Lhasa 850600, China

Received:2025-07-21 Accepted:2026-02-25 Online:2026-03-20 Published:2026-05-20
Contact: *ZHOU Ji-Qiong(jiqiong_zhou@sicau.edu.cn)
Supported by:
National Natural Science Foundation of China(32271776);Sichuan Provincial Natural Science Foundation(24NSFSC1813);Joint Fund Project of Science and Education in Sichuan Province(25LHJJ0196);Joint Fund Project of Science and Education in Sichuan Province(25LHJJ0190)

摘要/Abstract

摘要：

豆禾混播是修复西南退化草地及提升生产力的重要措施。丛枝菌根真菌(AMF)在其中发挥关键的养分调控作用。但AMF如何调控不同根系深度植物构建的混播群落氮吸收与分配机制仍不明晰。该研究根据植物的固氮特性和根系深度差异, 选取深根型豆科植物苜蓿(Medicago sativa)和浅根型豆科植物白三叶(Trifolium repens), 与两种禾本科植物鸭茅(Dactylis glomerata)和黑麦草(Lolium perenne)单播及混播, 设置不同的AMF接种处理(接种和不接种), 利用¹⁵N同位素示踪技术(分别标记3 cm浅层与25 cm深层土壤), 探究AMF对不同根系深度植物氮获取策略及群落功能的影响。结果表明, 混播显著提高了植物群落总生物量, 尤其以“白三叶+鸭茅+黑麦草”混播组合为最佳, 较白三叶或黑麦草单播分别显著增产54.78%和76.58%, 表明浅根型豆科植物与禾本科植物形成了显著的地下生态位互补, 增强了资源利用效率; 相反, 深根型豆科植物苜蓿与禾本科植物因根系空间布局重叠, 种间竞争加强, 混播优势不显著。接种AMF显著促进菌根依赖性较强的白三叶和苜蓿的生物量积累, 削弱了依赖性较弱的禾本科植物鸭茅和黑麦草的竞争优势, 从而提高了豆科植物在群落中的比例; 此外, AMF诱导了植物根系形态的可塑性, 显著增加了白三叶和黑麦草的根系长度。¹⁵N示踪结果发现, 两种混播较单播促进了豆科植物的固氮率, 接种AMF进一步强化了这一过程并降低了豆科植物对土壤¹⁵N的吸收。同时AMF缩小了禾本科植物在不同土层的¹⁵N吸收差异, 促进了其对深层土壤氮的高效利用。AMF通过增强豆科植物固氮效率和促进禾本科植物对深层土壤氮的获取, 实现了群落水平的资源优化配置。该研究验证“浅根型豆科+禾本科”混播并接种AMF是实现地下空间对接与地上高产的最佳策略, 为西南地区人工草地建植提供了理论借鉴。

关键词: 丛枝菌根真菌, 豆禾混播, 深根系, 浅根系, 氮吸收, 生物固氮, ¹⁵N示踪

Abstract:

Aims Legume-grass intercropping is an important strategy for restoring degraded grasslands and improving productivity in Southwest China. Arbuscular mycorrhizal fungi (AMF) play a key regulatory role in nutrient uptake within these systems. However, the mechanism by which AMF regulates nitrogen uptake and allocation in intercropping communities composed of plants with varying rooting depths remains unclear.

Methods Based on differences in nitrogen-fixing traits and root depths, this study selected a deep-rooted legume Medicago sativa and a shallow-rooted legume Trifolium repens, intercropped with two grass species, Dactylis glomerata and Lolium perenne. Monocultures and mixtures were established with or without AMF. Using ¹⁵N labeling at two soil depths 3 cm shallow and 25 cm deep, we explored the effects of AMF on the nitrogen acquisition strategies and community functions of plants with different root depths.

Important findings Intercropping significantly increased the total community of the plant community. The combination of T. repens + D. glomerata + L. perenne performed best, increasing yield by 54.78% and 76.58% compared to T. repens or L. perenne monocultures, respectively. This highlights a significant belowground niche complementarity between shallow-rooted legumes and grasses, which enhanced resource use efficiency. Conversely, the deep-rooted M. sativa showed spatial root overlap with grasses, intensifying interspecific competition and diminishing in insignificant intercropping advantages. AMF inoculation significantly promoted biomass accumulation in the strongly mycorrhizal-dependent legumes while moderating the competitive dominance of the less-dependent grasses, thereby increasing the legume proportion within the community. Furthermore, AMF induced plasticity in root morphology, significantly increasing the root length of T. repens and L. perenne.¹⁵N tracing revealed that intercropping promoted legume nitrogen fixation rates; AMF inoculation further strengthened this process and reduced the uptake of soil ¹⁵N by legumes. Meanwhile, AMF narrowed the difference in ¹⁵N uptake by grasses across soil layers, promoting more efficient utilization of deep-soil nitrogen. AMF achieved optimal resource allocation at the community level by enhancing legumes nitrogen fixation efficiency and promoting deep soil nitrogen acquisition by grasses. This study verifies that the shallow-rooted legume + grass mixture combined with AMF inoculation is the optimal strategy for achieving belowground spatial partitioning and high aboveground yield, providing a theoretical reference for the establishment of artificial grasslands in Southwest China.

Key words: arbuscular mycorrhizal fungi (AMF), legume-grass mixture, deep root system, shallow root system, nitrogen uptake, biological nitrogen fixation, ¹⁵N labeling

魏莉, 王鹏森, 刘珊, 樊锐, 黄楠, 张建国, 其美拉姆, 苟扬, 刘沫含, 黄婷, 周冀琼. 丛枝菌根真菌对豆禾混播植物垂直生态位养分吸收的影响. 植物生态学报, 2026, 50(3): 760-773. DOI: 10.17521/cjpe.2025.0272

WEI Li, WANG Peng-Sen, LIU Shan, FAN Rui, HUANG Nan, ZHANG Jian-Guo, Qimeilamu , GOU Yang, LIU Mo-Han, HUANG Ting, ZHOU Ji-Qiong. Arbuscular mycorrhizal fungi influence nutrient uptake along vertical niches in legume-grass mixtures. Chinese Journal of Plant Ecology, 2026, 50(3): 760-773. DOI: 10.17521/cjpe.2025.0272

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2025.0272

https://www.plant-ecology.com/CN/Y2026/V50/I3/760

图/表 10

图1 不同土层15N标记的实验设计示意图。AMF, 丛枝菌根真菌; -AMF, 未接种AMF; + AMF, 接种AMF。Dg, 鸭茅单播; Lp, 黑麦草单播; Ms, 苜蓿单播; Tr, 白三叶单播; Ms + Dg + Lp, 苜蓿+鸭茅+黑麦草混播; Tr + Dg + Lp, 白三叶+鸭茅+黑麦草混播。

Fig. 1 15N isotope labeling at various soil depths. AMF, arbuscular mycorrhizal fungi; -AMF, without AMF; +AMF, inoculation with AMF; Dg, Dactylis glomerata monoculture; Lp, Lolium perenne monoculture; Ms, Medicago sativa monoculture; Tr, Trifolium repens monoculture; Ms + Dg + Lp, Medicago sativa + Dactylis glomerata + Lolium perenne mix; Tr + Dg + Lp, Trifolium repens + Dactylis glomerata + Lolium perenne mix.

表1 种植方式(CS)、接种丛枝菌根真菌(AMF)及其交互作用对4种植物群落生物量影响的双因素方差分析

Table 1 Two-way ANOVA results of the effects of cropping system (CS), arbuscular mycorrhizal fungi inoculation (AMF), and their interaction on plant community biomass of four plant species

变量 Response variable		CS	AMF	CS × AMF
df		1/2	1	1/2
植物群落生物量 Plant community biomass
苜蓿 Medicago sativa	F	0.443	4.247	0.435
苜蓿 Medicago sativa	p	0.513	0.053	0.517
白三叶 Trifolium repens	F	67.118	9.526	1.301
白三叶 Trifolium repens	p	<0.001	0.006	0.267
鸭茅 Dactylis glomerata	F	4.940	3.113	0.212
鸭茅 Dactylis glomerata	p	0.014	0.088	0.810
黑麦草 Lolium perenne	F	20.072	1.601	0.624
黑麦草 Lolium perenne	p	<0.001	0.215	0.542

图2 种植方式和接种丛枝菌根真菌(AMF)对植物群落生物量(地上和地下总生物量)的影响。-AMF, 未接种AMF; +AMF, 接种AMF。Dg, 鸭茅单播; Lp, 黑麦草单播; Ms, 苜蓿单播; Tr, 白三叶单播; Ms + Dg + Lp, 苜蓿+鸭茅+黑麦草混播; Tr + Dg + Lp, 白三叶+鸭茅+黑麦草混播。不同小写字母表示处理间差异显著(p < 0.05)。

Fig. 2 Effects of culture system and inoculation with arbuscular mycorrhizal fungi (AMF) on plant community biomass (total aboveground and belowground biomass). -AMF, without AMF; +AMF, inoculation with AMF; Dg, Dactylis glomerata monoculture; Lp, Lolium perenne monoculture; Ms, Medicago sativa monoculture; Tr, Trifolium repens monoculture; Ms + Dg + Lp, Medicago sativa + Dactylis glomerata + Lolium perenne mix; Tr + Dg + Lp, Trifolium repens + Dactylis glomerata + Lolium perenne mix. Different lowercase letters denote significant differences between treatments (p < 0.05).

图3 种植方式和接种丛枝菌根真菌(AMF)对植物地上生物量和地下生物量的影响。-AMF, 未接种AMF; +AMF, 接种AMF。Dg, 鸭茅单播; Lp, 黑麦草单播; Ms, 苜蓿单播; Tr, 白三叶单播; Ms + Dg + Lp, 苜蓿+鸭茅+黑麦草混播; Tr + Dg + Lp, 白三叶+鸭茅+黑麦草混播。括号内的字母代表该植物在混播体系中的地上生物量和地下生物量。CS, 种植方式; AMF, 接种AMF处理; CS × AMF, 种植方式与接种AMF处理交互效应。每个图上方显示了双因素方差分析的结果。*, p < 0.05; **, p < 0.01; ***, p < 0.001。不同小写字母表示处理间差异显著(p < 0.05)。

Fig. 3 Effects of culture system and inoculation with arbuscular mycorrhizal fungi (AMF) on plant above biomass, below ground biomass. -AMF, without AMF; +AMF, inoculation with AMF; Dg, Dactylis glomerata monoculture; Lp, Lolium perenne monoculture; Ms, Medicago sativa monoculture; Tr, Trifolium repens monoculture; Ms + Dg + Lp, Medicago sativa + Dactylis glomerata + Lolium perenne mix; Tr + Dg + Lp, Trifolium repens + Dactylis glomerata + Lolium perenne mix. The letters in parentheses indicate the aboveground biomass or belowground biomass of that specific plant species with the nixed system. CS indicates the cropping system, AMF indicates AMF inoculation, and CS × AMF indicates their interaction effect. Two-way ANOVA results are shown above each panel: *, p < 0.05; ***, p < 0.001. Different lowercase letters indicate significant differences between treatments (p < 0.05).

图4 种植方式和丛枝菌根真菌接种(AMF)对植物根系长度的影响。-AMF, 未接种AMF; +AMF, 接种AMF; Dg, 鸭茅单播; Lp, 黑麦草单播; Ms, 苜蓿单播; Tr, 白三叶单播; Ms + Dg + Lp, 苜蓿+鸭茅+黑麦草混播; Tr + Dg + Lp, 白三叶+鸭茅+黑麦草混播。括号内的字母代表该植物在混插体系中的根系长度。CS, 种植方式; AMF, 接种AMF处理; CS × AMF, 种植方式与接种AMF处理交互效应。每个图上方显示了双因素方差分析的结果: *, p < 0.05; **, p < 0.01。不同小写字母表示处理间差异显著(p < 0.05)。

Fig. 4 Effects of culture system and inoculation with arbuscular mycorrhizal fungi (AMF) on plant root length. -AMF, without AMF; +AMF, inoculation with AMF; Dg, Dactylis glomerata monoculture; Lp, Lolium perenne monoculture; Ms, Medicago sativa monoculture; Tr, Trifolium repens monoculture; Ms + Dg + Lp, Medicago sativa + Dactylis glomerata + Lolium perenne mix; Tr + Dg + Lp, Trifolium repens + Dactylis glomerata + Lolium perenne mix. The letters in parentheses indicate the root length of that specific plant species within the mixed system. CS indicates the cropping system, AMF indicates AMF inoculation, and CS × AMF indicates their interaction effect. Two-way ANOVA results are shown above each panel: *, p < 0.05; **, p < 0.01; ***, p < 0.001. Different lowercase letters indicate significant differences between treatments (p < 0.05).

图5 种植方式对菌根侵染强度的影响。Dg, 鸭茅单播; Lp, 黑麦草单播; Ms, 苜蓿单播; Ms + Dg + Lp, 苜蓿+鸭茅+黑麦草混播; Tr, 白三叶单播; Tr + Dg + Lp, 白三叶+鸭茅+黑麦草混播。括号内的字母代表该植物在混播体系中的菌根侵染强度。不同小写字母表示处理间差异显著(p < 0.05)。

Fig. 5 Effects of culture systems on the intensity of mycorrhizal colonization. Dg, Dactylis glomerata monoculture; Lp, Lolium perenne monoculture; Ms, Medicago sativa monoculture; Ms + Dg + Lp, Medicago sativa + Dactylis glomerata + Lolium perenne mix; Tr, Trifolium repens monoculture; Tr + Dg + Lp, Trifolium repens + Dactylis glomerata + Lolium perenne mix. The letters in parentheses indicate the mycorrhizal colonization intensity of that specific plant species within the mixed system. Different lowercase letters indicate significant differences between treatments (p < 0.05).

图6 种植方式对地上植物生物量的菌根生长响应。Dg, 鸭茅单播; Lp, 黑麦草单播; Ms, 苜蓿单播; Ms + Dg + Lp, 苜蓿+鸭茅+黑麦草混播; Tr, 白三叶单播; Tr + Dg + Lp, 白三叶+鸭茅+黑麦草混播。括号内的字母代表该植物在混播体系中的菌根生长响应。不同小写字母表示处理间差异显著(p < 0.05)。

Fig. 6 Effect of culture system on the mycorrhizal growth response of plant aboveground biomass. Dg, Dactylis glomerata monoculture; Lp, Lolium perenne monoculture; Ms, Medicago sativa monoculture; Ms + Dg + Lp, Medicago sativa + Dactylis glomerata + Lolium perenne mix; Tr, Trifolium repens monoculture; Tr + Dg + Lp, Trifolium repens + Dactylis glomerata + Lolium perenne mix. The letters in parentheses indicate the mycorrhizal growth response of that specific plant species within the mixed system. Different lowercase letters indicate significant differences between treatments (p < 0.05).

图7 种植方式和接种丛枝菌根真菌(AMF)对豆科植物固氮率的影响。-AMF, 未接种AMF; +AMF, 接种AMF。Dg, 鸭茅单播; Lp, 黑麦草单播; Ms, 苜蓿单播; Ms + Dg + Lp, 苜蓿+鸭茅+黑麦草混播; Tr, 白三叶单播; Tr + Dg + Lp, 白三叶+鸭茅+黑麦草混播。括号内的字母代表该植物在混播体系中的固氮率。CS, 种植方式; AMF, 接种AMF处理; CS × AMF, 种植方式与接种AMF处理交互效应。25 cm, 25 cm处15N示踪剂标记深度; 3 cm, 3 cm处15N示踪剂标记深度。Depth, 15N示踪剂标记深度。每个图上方显示了三因素方差分析的结果: *, p < 0.05; **, p < 0.01; ***, p < 0.001。不同小写字母表示处理间差异显著(p < 0.05)。

Fig. 7 Effects of culture system and inoculation with arbuscular mycorrhizal fungi (AMF) on nitrogen fixation ratio of legumes (Ndfa). -AMF, without AMF; +AMF, inoculation with AMF; Dg, Dactylis glomerata monoculture; Lp, Lolium perenne monoculture; Ms, Medicago sativa monoculture; Ms + Dg + Lp, Medicago sativa + Dactylis glomerata + Lolium perenne mix; Tr, Trifolium repens monoculture; Tr + Dg + Lp, Trifolium repens + Dactylis glomerata + Lolium perenne mix. The letters in parentheses indicate the nitrogen fixation rate of that specific plant species within the mixed system. CS indicates cropping system, AMF indicates AMF inoculation treatment, and Depth indicates 15N tracer depth; CS × AMF indicates their interaction. 25 cm, 15N tracer labeling at 25 cm depth; 3 cm, 15N tracer labeling at 3 cm depth. Three-way ANOVA results are shown above each panel: *, p < 0.05; **, p < 0.01; ***, p < 0.001. Different lowercase letters indicate significant differences between treatments (p < 0.05).

图8 种植方式和接种丛枝菌根真菌(AMF)对地上部分植物氮含量的影响。-AMF, 未接种AMF; +AMF, 接种AMF。Dg, 鸭茅单播; Lp, 黑麦草单播; Ms, 苜蓿单播; Ms + Dg + Lp, 苜蓿+鸭茅+黑麦草混播; Tr, 白三叶单播; Tr + Dg + Lp, 白三叶+鸭茅+黑麦草混播。括号内的字母代表该植物在混播体系中的植物氮含量。CS, 种植方式; AMF, 接种AMF处理; CS × AMF, 种植方式与接种AMF处理交互效应: *, p < 0.05; **, p < 0.01; ***, p < 0.001。不同小写字母表示处理间差异显著(p < 0.05)。

Fig. 8 Effects of culture system and inoculation with arbuscular mycorrhizal fungi (AMF) on aboveground plant nitrogen content. -AMF, without AMF; +AMF, inoculation with AMF; Dg, Dactylis glomerata monoculture; Lp, Lolium perenne monoculture; Ms, Medicago sativa monoculture; Ms + Dg + Lp, Medicago sativa + Dactylis glomerata + Lolium perenne mix; Tr, Trifolium repens monoculture; Tr + Dg + Lp, Trifolium repens + Dactylis glomerata + Lolium perenne mix. The letters in parentheses indicate the plant nitrogen content of that specific plant species within the mixed system. CS indicates cultivation mode and AMF indicates inoculation mode of AMF, CS × AMF is the interaction effect of the two. Two-way ANOVA results are shown above each panel: *, p < 0.05; **, p < 0.01; ***, p < 0.001. Different lowercase letters indicate significant differences between treatments (p < 0.05).

图9 种植方式和接种丛枝菌根真菌(AMF)对植物示踪剂吸收量的影响。-AMF, 未接种AMF; +AMF, 接种AMF。Dg, 鸭茅单播; Lp, 黑麦草单播; Ms, 苜蓿单播; Ms + Dg + Lp, 苜蓿+鸭茅+黑麦草混播; Tr, 白三叶单播; Tr + Dg + Lp, 白三叶+鸭茅+黑麦草混播。括号内的字母代表该植物在混播体系中的植物示踪剂吸收量。CS, 种植方式; AMF, 接种AMF处理; CS × AMF, 种植方式与接种AMF处理交互效应。25 cm, 25 cm处15N示踪剂标记深度; 3 cm, 3 cm处15N示踪剂标记深度。Depth, 15N示踪剂标记深度。每个图上方显示了双因素方差分析的结果: *, p < 0.05; **, p < 0.01; ***, p < 0.001。不同小写字母表示处理间差异显著(p < 0.05)。

Fig. 9 Effects of culture system and inoculation with arbuscular mycorrhizal fungi (AMF) on plant uptake of tracer. -AMF, without AMF; +AMF, inoculation with AMF; Dg, Dactylis glomerata monoculture; Lp, Lolium perenne monoculture; Ms, Medicago sativa monoculture; Ms + Dg + Lp, Medicago sativa + Dactylis glomerata + Lolium perenne mix; Tr, Trifolium repens monoculture; Tr + Dg + Lp, Trifolium repens + Dactylis glomerata + Lolium perenne mix. The letters in parentheses indicate the plant tracer uptake of that specific plant species within the mixed system. CS indicates cultivation mode and AMF indicates inoculation mode of AMF, CS × AMF is the interaction effect of the two. Depth: depth of 15N tracer labeling; 25 cm, 15N tracer labeling at 25 cm depth; 3 cm, 15N tracer labeling at 3 cm depth: *, p < 0.05; **, p < 0.01; ***, p < 0.001. Different lowercase letters indicate significant differences between treatments (p < 0.05).

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丛枝菌根真菌对豆禾混播植物垂直生态位养分吸收的影响

Arbuscular mycorrhizal fungi influence nutrient uptake along vertical niches in legume-grass mixtures

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