Research progress on root exudates and rhizosphere effects of tree species associated with different mycorrhizal types

doi:10.17521/cjpe.2024.0196

Abstract

Abstract:

Changes in belowground ecological processes in the context of global change have become one of the hot issues of concern in the field of ecology. Mycorrhiza are symbiotic relationships between plant roots and mycorrhizal fungi, widely distributed in terrestrial ecosystems. As two major types of mycorrhizal fungi related to trees, arbuscular mycorrhizal fungi and ectomycorrhizal fungi exhibit significant differences in morphology and function. Root exudates, as an important medium for exchange of matter, energy, and information between plants and soil, play a crucial role in soil carbon dynamics. The root exudates of tree species associated with different mycorrhizal types can actively respond to environmental changes by continuously adjusting their quantity and chemical composition, and influence the belowground carbon dynamics and cycling processes of forest ecosystems. Currently, the composition and function of root exudates from different types of mycorrhizal fungi, as well as the variation and impact on plant and soil, remain unclear. Therefore, this article combines current frontiers in the field both domestically and internationally, and summarizes the root exudate characteristics, influencing mechanisms, and rhizosphere effects of tree species associated with different mycorrhizal types. This review is expected to provide references for further research on the response and adaptation mechanism of root systems and exudates to global changes. In addition, it also proposes directions for future research on root exudates among different types of mycorrhizal fungi that require further investigation: (1) strengthening systematic research on root exudates among different types of mycorrhizal fungi; (2) studying the influence mechanism of mycorrhizal type on root exudates in combination with other environmental factors; (3) using more precise technological means to comprehensively understand the changes in root exudate characteristics among different types of mycorrhizal fungi; (4) deeply revealing the influencing mechanism of root exudates among different types of mycorrhizal fungi from the perspective of plant physiology and metabolism; (5) conducting long-term dynamic monitoring and simulation experiments on different types of mycorrhizal fungi to predict their impact on soil ecological processes.

Key words: root exudates, arbuscular mycorrhiza, ectomycorrhiza, below-ground carbon allocation, impact mechanism, rhizosphere effect

LIANG Tian-Hao, XIONG De-Cheng, LIU Yuan-Hao, DU Xu-Long, YANG Zhi-Jie, HUANG Jin-Xue. Research progress on root exudates and rhizosphere effects of tree species associated with different mycorrhizal types[J]. Chin J Plant Ecol, 2025, 49(7): 1038-1052.

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研究区域 Study area	研究对象 Study object	研究类型 Type	测定方法 Method	速率(R)特征 Rate (R) characteristic	影响机理 Impact mechanism	参考文献 Reference
亚热带 Subtropical zone	10种丛枝菌根树种和10种外生菌根树种 10 AM species and 10 ECM species	野外实验 Field experiment
温带 Temperate zone	黄桦、糖槭 Betula allegheniensis, Acer saccharum	盆栽实验 Pot experiment	土培收集: 收集粘附在细根周围的根际土, 通过振荡离心或过滤得到根系分泌物。由于需要通过淋洗将根系与土壤进行分离, 难以完全排除土粒本身的影响, 一定程度上造成根系损伤, 收集到的根系分泌物可能含有根系本身的分泌物质 Soil cultivation collection: collecting the rhizosphere soil surrounding the adhering fine roots, and obtain the root exudates by shock centrifugation or filtration. Due to the need to separate the root from the soil through washing, it is difficult to completely exclude the influence of the soil particles themselves, causing root damage to some extent, and the collected root exudates may contain exudation material from the root itself	R_ECM = 2R_AM	外生菌根树种比丛枝菌根树种通常可以分配更多的碳水化合物到根系分泌物中 ECM tree species can usually allocate more carbohydrates to root exudates than AM tree species	Phillips & Fahey, 2006
	北美白橡木、北美水青冈、糖槭和北美鹅掌楸 Quercus alba, Fagus grandifolia, Acer saccharum and Liriodendron tulipifera	野外实验 Field experiment	参考Phillips等(2008)的野外原位收集方法 Refer to the field in-situ collection method in Phillips et al. (2008)	R_ECM = 2R_AM	外生菌根树种以有机养分循环过程为主, 通过分泌更多的碳(C)以促进土壤有机质(SOM)分解, 获取养分 ECM species are dominated by the organic nutrient cycling process and secrete more carbon (C) to obtain nutrients from SOM	Yin et al., 2014
	4种丛枝菌根树种和3种外生菌根树种 4 AM tree species and 3 ECM tree species	野外实验 Field experiment	参考Phillips等(2008)的野外原位收集方法 Refer to the field in-situ collection method in Phillips et al. (2008)	R_ECM > R_AM	外生菌根的根外菌丝网络数量级远高于AM菌根, 显著促进了ECM树种根系的分泌 The extrarhizal hyphal network of ECM mycorrhizae is an order of magnitude higher than that of AM mycorrhizae, which significantly promotes the root exudation of ECM tree species	Jiang et al., 2021
温带 Temperate zone	枹栎、具柄冬青 Quercus serrata, Ilex pedunculosa	野外实验 Field experiment	参考Phillips等(2008)的野外原位收集方法 Refer to the field in-situ collection method in Phillips et al. (2008)	R_AM = 2R_ECM	根系分泌速率与叶氮(N)含量正相关, 这是因为根系分泌速率与分配给根系的活性C含量有关, 植物将C分配到根系分泌物中以获取N可能是一种有效的促进生长的策略 Root exudation rate is positively correlated with leaf nitrogen (N) content because root exudation rate is related to the active C content assigned to roots, and plants assigning C to root exudates to obtain N may be an effective growth-promoting strategy	Sun et al., 2017
	6种丛枝菌根树种和8种外生菌根树种 6 AM species and 8 ECM species	野外实验 Field experiment	参考Phillips等(2008)的野外原位收集方法 Refer to the field in-situ collection method in Phillips et al. (2008)	R_ECM ≈ R_AM	不同物种土壤有效氮的空间异质性较大 Large spatial heterogeneity of soil available N under different species	Han et al., 2020
	4种丛枝菌根树种和4种外生菌根树种 4 AM species and 4 ECM species	盆栽实验 Pot experiment	土培收集 Soil incubation collection	R_ECM = 2R_AM	外生菌根树木比丛枝菌根树木具有更强的抗旱能力以及更高的导水效率和养分利用效率, 这就导致在干旱胁迫下AM树种可能比ECM树种根系分泌量更大 ECM trees have better drought resistance and higher water diversion efficiency than AM trees, which leads to greater root exudation in AM tree species than ECM tree species under drought stress	Liese et al., 2018

研究区域 Study area	研究对象 Study object	研究类型 Type	测定方法 Method	速率(R)特征 Rate (R) characteristic	影响机理 Impact mechanism	参考文献 Reference
亚热带 Subtropical zone	10种丛枝菌根树种和10种外生菌根树种 10 AM species and 10 ECM species	野外实验 Field experiment
温带 Temperate zone	黄桦、糖槭 Betula allegheniensis, Acer saccharum	盆栽实验 Pot experiment	土培收集: 收集粘附在细根周围的根际土, 通过振荡离心或过滤得到根系分泌物。由于需要通过淋洗将根系与土壤进行分离, 难以完全排除土粒本身的影响, 一定程度上造成根系损伤, 收集到的根系分泌物可能含有根系本身的分泌物质 Soil cultivation collection: collecting the rhizosphere soil surrounding the adhering fine roots, and obtain the root exudates by shock centrifugation or filtration. Due to the need to separate the root from the soil through washing, it is difficult to completely exclude the influence of the soil particles themselves, causing root damage to some extent, and the collected root exudates may contain exudation material from the root itself	R_ECM = 2R_AM	外生菌根树种比丛枝菌根树种通常可以分配更多的碳水化合物到根系分泌物中 ECM tree species can usually allocate more carbohydrates to root exudates than AM tree species	Phillips & Fahey, 2006
	北美白橡木、北美水青冈、糖槭和北美鹅掌楸 Quercus alba, Fagus grandifolia, Acer saccharum and Liriodendron tulipifera	野外实验 Field experiment	参考Phillips等(2008)的野外原位收集方法 Refer to the field in-situ collection method in Phillips et al. (2008)	R_ECM = 2R_AM	外生菌根树种以有机养分循环过程为主, 通过分泌更多的碳(C)以促进土壤有机质(SOM)分解, 获取养分 ECM species are dominated by the organic nutrient cycling process and secrete more carbon (C) to obtain nutrients from SOM	Yin et al., 2014
	4种丛枝菌根树种和3种外生菌根树种 4 AM tree species and 3 ECM tree species	野外实验 Field experiment	参考Phillips等(2008)的野外原位收集方法 Refer to the field in-situ collection method in Phillips et al. (2008)	R_ECM > R_AM	外生菌根的根外菌丝网络数量级远高于AM菌根, 显著促进了ECM树种根系的分泌 The extrarhizal hyphal network of ECM mycorrhizae is an order of magnitude higher than that of AM mycorrhizae, which significantly promotes the root exudation of ECM tree species	Jiang et al., 2021
温带 Temperate zone	枹栎、具柄冬青 Quercus serrata, Ilex pedunculosa	野外实验 Field experiment	参考Phillips等(2008)的野外原位收集方法 Refer to the field in-situ collection method in Phillips et al. (2008)	R_AM = 2R_ECM	根系分泌速率与叶氮(N)含量正相关, 这是因为根系分泌速率与分配给根系的活性C含量有关, 植物将C分配到根系分泌物中以获取N可能是一种有效的促进生长的策略 Root exudation rate is positively correlated with leaf nitrogen (N) content because root exudation rate is related to the active C content assigned to roots, and plants assigning C to root exudates to obtain N may be an effective growth-promoting strategy	Sun et al., 2017
	6种丛枝菌根树种和8种外生菌根树种 6 AM species and 8 ECM species	野外实验 Field experiment	参考Phillips等(2008)的野外原位收集方法 Refer to the field in-situ collection method in Phillips et al. (2008)	R_ECM ≈ R_AM	不同物种土壤有效氮的空间异质性较大 Large spatial heterogeneity of soil available N under different species	Han et al., 2020
	4种丛枝菌根树种和4种外生菌根树种 4 AM species and 4 ECM species	盆栽实验 Pot experiment	土培收集 Soil incubation collection	R_ECM = 2R_AM	外生菌根树木比丛枝菌根树木具有更强的抗旱能力以及更高的导水效率和养分利用效率, 这就导致在干旱胁迫下AM树种可能比ECM树种根系分泌量更大 ECM trees have better drought resistance and higher water diversion efficiency than AM trees, which leads to greater root exudation in AM tree species than ECM tree species under drought stress	Liese et al., 2018

研究区域 Study area	研究对象 Study object	研究类型 Study type	测定方法 Method	化学组成特征 Chemical composition characteristic	影响机理 Impact mechanism	参考文献 Reference
亚热带 Subtropical zone	10种丛枝菌根树种和10种外生菌根树种 10 AM species and 10 ECM species	野外实验 Field experiment	参考Phillips等(2008)的野外原位收集方法 Refer to the field in-situ collection method in Phillips et al. (2008)	有机酸(ECM树种比AM树种分泌多) Organic acids (ECM species secrete more than AM species)	磷(P)限制下, 外生菌根树种根系分泌大量的有机酸, 以便释放出土壤中顽固的有机磷和矿质磷 Under phosphorous (P) deficiency, roots of ECM tree species secrete large amounts of organic acids to facilitate the release of recalcitrant organophosphorus and mineral phosphorus from the soil	Jiang et al., 2022
亚热带 Subtropical zone	10种丛枝菌根树种和10种外生菌根树种 10 AM species and 10 ECM species	野外实验 Field experiment	参考Phillips等(2008)的野外原位收集方法 Refer to the field in-situ collection method in Phillips et al. (2008)	ACP (丛枝菌根树种比外生菌根树种分泌更多) ACP (AM species secrete more than ECM species)	丛枝菌根树种根系的形态和分泌物比外生菌根树种更容易受到无机营养缺乏的影响 The root morphology and exudates of AM species are more susceptible to inorganic nutrient deficiency than those of ECM species	Jiang et al., 2022
温带 Temperate zone	5种丛枝菌根树种和6种外生菌根树种 5 AM species and 6 ECM species	野外实验 Field experiment	土培收集 Soil incubation collection	碳水化合物(ECM树种分泌多, AM树种分泌少) Carbohydrates (high in ECM species, low in AM species)	外生菌根真菌通常比丛枝菌根真菌具有更高的生物量 ECM fungi usually have a higher biomass than AM fungi	Yin et al., 2014
	美国白梣、糖槭、北美水青冈和加拿大铁杉 Fraxinus americana, Acer saccharum, Fagus grandifolia and Tsuga canadensis	野外实验 Field experiment	参考Phillips等(2008)的野外原位收集方法 Refer to the field in-situ collection method in Phillips et al. (2008)	土壤胞外酶(外生菌根树种比丛枝菌根树种分泌多) Soil extracellular enzymes (ECM species secrete more than AM species)	外生菌根树种比丛枝菌根树种分泌较多的活性C, 促进土壤微生物合成较多的胞外酶 ECM species secrete more active C than AM species, promoting the synthesis of more extracellular enzymes by soil microorganisms	Phillips et al., 2013
	木荷、冬青栎 Schima superba, Quercus ilex	野外试验 Field experiment	参考Phillips等(2008)的野外原位收集方法 Refer to the field in-situ collection method in Phillips et al. (2008)	干旱条件下, 外生菌根树种糖类、氨基酸分泌增加, 有机酸减少; 丛枝菌根树种糖类、氨基酸分泌减少, 有机酸增加 Under drought conditions, the exudation of sugars and amino acids in ECM species increased while organic acid decreased; the secretion of sugars and organic acids in AM species decreased	干旱下, 根系分泌速率与树种生长速度正相关, 而外生菌根树种比丛枝菌根树种生长速度快 Under drought, the rate of root exudates was positively correlated with the growth rate of the tree species, while ECM species grew faster than AM species	Jiang et al., 2022

研究区域 Study area	研究对象 Study object	研究类型 Study type	测定方法 Method	化学组成特征 Chemical composition characteristic	影响机理 Impact mechanism	参考文献 Reference
亚热带 Subtropical zone	10种丛枝菌根树种和10种外生菌根树种 10 AM species and 10 ECM species	野外实验 Field experiment	参考Phillips等(2008)的野外原位收集方法 Refer to the field in-situ collection method in Phillips et al. (2008)	有机酸(ECM树种比AM树种分泌多) Organic acids (ECM species secrete more than AM species)	磷(P)限制下, 外生菌根树种根系分泌大量的有机酸, 以便释放出土壤中顽固的有机磷和矿质磷 Under phosphorous (P) deficiency, roots of ECM tree species secrete large amounts of organic acids to facilitate the release of recalcitrant organophosphorus and mineral phosphorus from the soil	Jiang et al., 2022
亚热带 Subtropical zone	10种丛枝菌根树种和10种外生菌根树种 10 AM species and 10 ECM species	野外实验 Field experiment	参考Phillips等(2008)的野外原位收集方法 Refer to the field in-situ collection method in Phillips et al. (2008)	ACP (丛枝菌根树种比外生菌根树种分泌更多) ACP (AM species secrete more than ECM species)	丛枝菌根树种根系的形态和分泌物比外生菌根树种更容易受到无机营养缺乏的影响 The root morphology and exudates of AM species are more susceptible to inorganic nutrient deficiency than those of ECM species	Jiang et al., 2022
温带 Temperate zone	5种丛枝菌根树种和6种外生菌根树种 5 AM species and 6 ECM species	野外实验 Field experiment	土培收集 Soil incubation collection	碳水化合物(ECM树种分泌多, AM树种分泌少) Carbohydrates (high in ECM species, low in AM species)	外生菌根真菌通常比丛枝菌根真菌具有更高的生物量 ECM fungi usually have a higher biomass than AM fungi	Yin et al., 2014
	美国白梣、糖槭、北美水青冈和加拿大铁杉 Fraxinus americana, Acer saccharum, Fagus grandifolia and Tsuga canadensis	野外实验 Field experiment	参考Phillips等(2008)的野外原位收集方法 Refer to the field in-situ collection method in Phillips et al. (2008)	土壤胞外酶(外生菌根树种比丛枝菌根树种分泌多) Soil extracellular enzymes (ECM species secrete more than AM species)	外生菌根树种比丛枝菌根树种分泌较多的活性C, 促进土壤微生物合成较多的胞外酶 ECM species secrete more active C than AM species, promoting the synthesis of more extracellular enzymes by soil microorganisms	Phillips et al., 2013
	木荷、冬青栎 Schima superba, Quercus ilex	野外试验 Field experiment	参考Phillips等(2008)的野外原位收集方法 Refer to the field in-situ collection method in Phillips et al. (2008)	干旱条件下, 外生菌根树种糖类、氨基酸分泌增加, 有机酸减少; 丛枝菌根树种糖类、氨基酸分泌减少, 有机酸增加 Under drought conditions, the exudation of sugars and amino acids in ECM species increased while organic acid decreased; the secretion of sugars and organic acids in AM species decreased	干旱下, 根系分泌速率与树种生长速度正相关, 而外生菌根树种比丛枝菌根树种生长速度快 Under drought, the rate of root exudates was positively correlated with the growth rate of the tree species, while ECM species grew faster than AM species	Jiang et al., 2022

环境因素 Environmental factor	菌根类型 Mycorrhizal type	对树种根系分泌物的影响 Effect on root exudates of tree species		参考文献 Reference
环境因素 Environmental factor	菌根类型 Mycorrhizal type	方向 Direction	程度 Level	参考文献 Reference
CO₂浓度升高 Elevation of the CO₂ concentration	丛枝菌根 AM	正向 Positive	显著 Significant	Yang et al., 2023
CO₂浓度升高 Elevation of the CO₂ concentration	外生菌根 ECM	正向 Positive	不显著 Not significant	Yang et al., 2023
氮沉降 Nitrogen deposition	丛枝菌根 AM	正向 Positive	显著 Significant	Midgley & Phillips, 2016
氮沉降 Nitrogen deposition	外生菌根 ECM	正向 Positive	不显著 Not significant	Midgley & Phillips, 2016
干旱 Drought	丛枝菌根 AM	正向 Positive	不显著 Not significant	Mohan et al., 2014
干旱 Drought	外生菌根 ECM	正向 Positive	显著 Significant	Mohan et al., 2014