外生菌根真菌: 高寒生态系统碳氮循环的关键驱动者

doi:10.17521/cjpe.2025.0350

植物生态学报 ›› 2026, Vol. 50 ›› Issue (3): 584-599.DOI: 10.17521/cjpe.2025.0350 cstr: 32100.14.cjpe.2025.0350

外生菌根真菌: 高寒生态系统碳氮循环的关键驱动者

谭秋颜¹^,², 张清¹^,³, 高程⁴, 褚海燕¹^,², 杨腾¹^,²^,⁵^,^*()

¹ 中国科学院南京土壤研究所土壤与农业可持续发展全国重点实验室, 南京 211135
² 中国科学院大学, 北京 100049
³ 南京信息工程大学, 南京 210044
⁴ 中国科学院微生物研究所微生物多样性与资源创新利用全国重点实验室, 北京 100101
⁵ 中国科学院大学南京学院, 南京 211135

收稿日期:2025-09-24 接受日期:2026-02-25 出版日期:2026-03-20 发布日期:2026-05-18
通讯作者: *杨腾(tyang@issas.ac.cn)
基金资助:
国家自然科学基金(42277308);中国科学院青年创新促进会项目(2022315)

Ectomycorrhizal fungi: key drivers of carbon and nitrogen cycling in alpine ecosystems

TAN Qiu-Yan¹^,², ZHANG Qing¹^,³, GAO Cheng⁴, CHU Hai-Yan¹^,², YANG Teng¹^,²^,⁵^,^*()

¹ State Key Laboratory of Soil and Sustainable Agriculture, Institute of Soil Science, Chinese Academy of Sciences, Nanjing 211135, China
² University of Chinese Academy of Sciences, Beijing 100049, China
³ Nanjing University of Information Science and Technology, Nanjing 210044, China
⁴ State Key Laboratory of Microbial Diversity and Innovative Utilization of Microbial Resources, Institute of Microbiology, Chinese Academy of Sciences, Beijing 100101, China
⁵ and University of Chinese Academy of Sciences, Nanjing, Nanjing 211135, China

Received:2025-09-24 Accepted:2026-02-25 Online:2026-03-20 Published:2026-05-18
Contact: *YANG Teng(tyang@issas.ac.cn)
Supported by:
National Natural Science Foundation of China(42277308);Youth Innovation Promotion Association of the Chinese Academy of Sciences(2022315)

摘要/Abstract

摘要：

高寒生态系统是高纬度、高海拔、气候寒冷、冻土广泛分布的地区, 也是全球变化的敏感区。外生菌根真菌是高寒生态系统优势植物的主要共生微生物类群, 通过独特的外生菌根结构与宿主植物建立互惠关系, 促进幼苗生长, 提升植物对水分和养分的吸收, 增强植物对生物和非生物胁迫的抗性。外生菌根真菌通过“碳输入-固持”和“氮矿化-吸收”双路径调控高寒生态系统的物质循环过程, 是高寒生态系统碳氮循环的关键驱动者。该文综述了外生菌根真菌在高寒生态系统碳氮循环方面的关键作用和全球变化下的响应特征, 强调了在全球变化背景下, 保护外生菌根真菌多样性及其关键类群以维持高寒生态系统碳氮循环过程和生态系统稳定的重要价值, 为高寒生态系统脆弱生境的保护和应对全球变化威胁提供了科学参考。未来需进一步加强外生菌根真菌功能性状与多样性特征对多重全球变化因子的响应与反馈调节机制研究, 并将菌根生态学的研究纳入“One Health”框架, 更好地服务于生态系统与人类健康。

关键词: 外生菌根真菌, 高寒生态系统, 碳循环, 氮循环, 全球变化

Abstract:

Alpine ecosystems, characterized by high latitudes, high altitudes, cold climates, and extensive permafrost, are highly sensitive to global climate change. Ectomycorrhizal fungi (ECMF) represent a major group of symbiotic microorganisms associated with dominant plant species in these ecosystems. By forming unique ectomycorrhizal structures with host plants, these fungi establish mutualistic relationships that promote seedling growth, enhance water and nutrient uptake, and improve host resistance to biotic and abiotic stresses. Through dual pathways of “carbon input-stabilization” and “nitrogen mineralization-uptake”, ECMF regulate material cycling and serve as key drivers of carbon and nitrogen dynamics. This review summarizes the critical roles of ECMF in carbon and nitrogen cycling within alpine ecosystems and their responses to climate change. It emphasizes the importance of preserving ECMF diversity and key functional groups to maintain ecosystem stability and nutrient processing under global change scenarios. Such conservation efforts provide essential scientific support for protecting vulnerable alpine habitats and mitigating climate-related threats. Future research should further investigate the response mechanisms and feedback regulations of ECMF functional traits and diversity under multiple-factor climate change. Integrating mycorrhizal ecology into the “One Health” framework will better serve the holistic health of both ecosystem and human societies.

Key words: ectomycorrhizal fungi, alpine ecosystem, carbon cycle, nitrogen cycle, climate change

谭秋颜, 张清, 高程, 褚海燕, 杨腾. 外生菌根真菌: 高寒生态系统碳氮循环的关键驱动者. 植物生态学报, 2026, 50(3): 584-599. DOI: 10.17521/cjpe.2025.0350

TAN Qiu-Yan, ZHANG Qing, GAO Cheng, CHU Hai-Yan, YANG Teng. Ectomycorrhizal fungi: key drivers of carbon and nitrogen cycling in alpine ecosystems. Chinese Journal of Plant Ecology, 2026, 50(3): 584-599. DOI: 10.17521/cjpe.2025.0350

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2025.0350

https://www.plant-ecology.com/CN/Y2026/V50/I3/584

图/表 3

图1 外生菌根真菌(ECMF)在高寒生态系统碳氮循环中的作用。A, 外生菌根真菌在碳循环中的作用。①光合碳通过外生菌根真菌输入土壤有机碳库; ②外生菌根真菌通过提高植物光合效率, 促进生态系统碳输入; ③加吉尔效应: 外生菌根真菌与腐生菌竞争氮, 抑制凋落物分解; ④外生菌根真菌菌丝残体、渗出物和代谢物输入土壤有机碳库(续埋效应); ⑤外生菌根真菌菌丝残体和渗出物转化为难分解的矿物结合态有机碳(MAOC), 稳定土壤有机碳库; ⑥土壤有机碳库通过酶促分解等作用被分解。绿色箭头代表碳的移动, 褐色箭头代表真菌的作用。B, 外生菌根真菌在氮循环中的作用。①外生菌根真菌分泌的蛋白酶和几丁质酶分解大量土壤有机氮为氨基酸和几丁质氮, 供植物吸收; ②植物直接通过根系吸收少量有机氮; ③大量无机氮经根系供给植物; ④植物通过外生菌根真菌吸收小分子有机氮; ⑤少量无机氮经外生菌根真菌的菌丝体供给植物; ⑥氨基酸和几丁质氮进一步矿化为无机氮。红色和蓝色分别代表有机氮和无机氮的移动, 实线代表经过外生菌根真菌的氮移动途径, 虚线代表不经过外生菌根真菌的氮移动途径。

Fig. 1 Role of ectomycorrhizal fungi (ECMF) in carbon and nitrogen cycling in alpine ecosystems. A, Role of ECMF in carbon cycling. ① Input of photosynthetic carbon into the soil organic carbon pool via ECMF; ② ECMF enhance ecosystem carbon input by improving plant photosynthetic efficiency; ③ Gadgil effect: ECMF compete with saprotrophic fungi for nitrogen, thereby suppressing litter decomposition; ④ Input of ECMF mycelia necromass, exudates, and metabolites into the soil organic carbon pool (entombing effect); ⑤ Transformation of ECMF mycelia necromass and exudates into mineral-associated organic carbon (MAOC), which stabilizes the soil organic carbon pool; ⑥ Soil organic carbon pools are decomposed through enzymatic decomposition and other processes. Green arrows represent carbon flow; brown arrows represent fungal-mediated processes. MCP: Microbial carbon pump. B, Role of ECMF in nitrogen cycling. ① Proteases and chitinases secreted by ECMF decompose soil organic nitrogen into amino acids and chitin-derived nitrogen for plant uptake; ② Plants directly absorb a small amount of organic nitrogen via roots; ③ A large amount of inorganic nitrogen is supplied to plants through root uptake; ④ Plants absorb low-molecular-weight organic nitrogen via ECMF; ⑤ A small amount of inorganic nitrogen is supplied to plants through ECMF mycelia; ⑥ Amino acids and chitin-derived nitrogen are further mineralized into inorganic nitrogen. Red and blue arrows represent organic and inorganic nitrogen flow, respectively; solid lines indicate ECMF-mediated pathways, and dashed lines indicate pathways not involving ECMF.

表1 全球变化因子对高寒生态系统外生菌根真菌驱动的碳氮循环过程的影响

Table 1 Impacts of climate change factors on carbon and nitrogen cycling processes driven by ectomycorrhizal fungi (ECMF) in alpine ecosystems

全球变化因子 Global change factor	生态系统/地点 Ecosystem/Location	碳氮循环过程变化 Changes in C/N cycling processes	参考文献 Reference
全球变暖 Global warming	北方森林 Boreal forest	限制地下外生菌根网络碳氮养分通道的形成和发展 Restrict the formation and development of underground ectomycorrhizal network pathways for carbon and nitrogen transfer	Fernandez et al., 2023
全球变暖 Global warming	北极苔原 Arctic tundra	增加外生菌根真菌菌丝体生物量, 更多光合碳经外生菌根真菌进入土壤; 菌丝体氮含量提高, 外生菌根真菌介导的氮矿化过程加强 Increase mycelial biomass of ECMF, and more photosynthetic carbon entering the soil via ECMF; enhance nitrogen content in mycelia and strengthen the ECMF-mediated nitrogen mineralization process	Clemmensen et al., 2006; Deslippe et al., 2011; Morgado et al., 2015
氮沉降 Nitrogen deposition	北方森林 Boreal forest	减少光合碳流向外生菌根真菌; 减少外生菌根真菌对有机氮的矿化 Reduce the flow of photosynthetic carbon to ECMF; reduce the mineralization of organic nitrogen by ECMF	Högberg et al., 2010; Jörgensen et al., 2024
CO₂浓度升高 Elevated CO₂concentration	北方森林 Boreal forest	增加外生菌根真菌残体碳的积累, 促进土壤碳固持; 增加外生菌根真菌的氮获取能力, 促进植物的氮吸收 Increase the accumulation of necromass carbon from ECMF, promoting soil carbon sequestration; enhance the nitrogen acquisition capacity of ECMF, facilitating plant nitrogen uptake	Liu et al., 2024; Reay et al., 2025
干旱 Drought	北方森林 Boreal forest	减少外生菌根真菌的定殖, 减弱加吉尔效应, 加速土壤有机质分解 Reduce colonization by ECMF, weaken the Gadgil effect, and therefore accelerate the decomposition of soil organic matter	Kilpeläinen et al., 2017; Nieves et al., 2024; Koelemeijer et al., 2025

表1 全球变化因子对高寒生态系统外生菌根真菌驱动的碳氮循环过程的影响

Table 1 Impacts of climate change factors on carbon and nitrogen cycling processes driven by ectomycorrhizal fungi (ECMF) in alpine ecosystems

全球变化因子 Global change factor	生态系统/地点 Ecosystem/Location	碳氮循环过程变化 Changes in C/N cycling processes	参考文献 Reference
全球变暖 Global warming	北方森林 Boreal forest	限制地下外生菌根网络碳氮养分通道的形成和发展 Restrict the formation and development of underground ectomycorrhizal network pathways for carbon and nitrogen transfer	Fernandez et al., 2023
全球变暖 Global warming	北极苔原 Arctic tundra	增加外生菌根真菌菌丝体生物量, 更多光合碳经外生菌根真菌进入土壤; 菌丝体氮含量提高, 外生菌根真菌介导的氮矿化过程加强 Increase mycelial biomass of ECMF, and more photosynthetic carbon entering the soil via ECMF; enhance nitrogen content in mycelia and strengthen the ECMF-mediated nitrogen mineralization process	Clemmensen et al., 2006; Deslippe et al., 2011; Morgado et al., 2015
氮沉降 Nitrogen deposition	北方森林 Boreal forest	减少光合碳流向外生菌根真菌; 减少外生菌根真菌对有机氮的矿化 Reduce the flow of photosynthetic carbon to ECMF; reduce the mineralization of organic nitrogen by ECMF	Högberg et al., 2010; Jörgensen et al., 2024
CO₂浓度升高 Elevated CO₂concentration	北方森林 Boreal forest	增加外生菌根真菌残体碳的积累, 促进土壤碳固持; 增加外生菌根真菌的氮获取能力, 促进植物的氮吸收 Increase the accumulation of necromass carbon from ECMF, promoting soil carbon sequestration; enhance the nitrogen acquisition capacity of ECMF, facilitating plant nitrogen uptake	Liu et al., 2024; Reay et al., 2025
干旱 Drought	北方森林 Boreal forest	减少外生菌根真菌的定殖, 减弱加吉尔效应, 加速土壤有机质分解 Reduce colonization by ECMF, weaken the Gadgil effect, and therefore accelerate the decomposition of soil organic matter	Kilpeläinen et al., 2017; Nieves et al., 2024; Koelemeijer et al., 2025

图2 外生菌根真菌(ECMF)促进土壤健康、森林健康、食品健康、人类健康。外生菌根真菌通过与植物根系形成共生关系, 增加土壤碳固持、稳定土壤结构, 在树木促生、抗病等方面发挥关键作用, 促进土壤和森林健康。健康的土壤可以培育健康的植物, 提供健康的食品, 促进人类健康; 健康的森林通过提供多种生态系统服务进一步促进人类健康。外生菌根真菌子实体(如松茸、松露、牛肝菌等)还具有食用和药用的价值。

Fig. 2 Ectomycorrhizal fungi (ECMF) function as vital links between soil, forest, food, and human health. By forming symbiotic relationships with plant roots, ECMF enhance soil carbon sequestration and stabilize soil structure, while also playing key roles in tree growth promotion and disease resistance, thereby contributing to soil and forest health. Healthy soil supports the growth of healthy plants, which provide nutritious food and promote human health; healthy forests further enhance human well-being by delivering diverse ecosystem services. Additionally, the fruiting bodies of ECMF (such as matsutake, truffles, and porcini) possess edible and medicinal value.

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外生菌根真菌: 高寒生态系统碳氮循环的关键驱动者

Ectomycorrhizal fungi: key drivers of carbon and nitrogen cycling in alpine ecosystems

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