香柱菌属内生真菌影响宿主禾草微生物群落结构与多样性的研究进展

doi:10.11686/cyxb2025402

摘要/Abstract

摘要：

香柱菌属内生真菌主要定殖于宿主禾草的地上组织，并与宿主形成稳定的非致病性共生体系。该类真菌不仅增强宿主对生物和非生物胁迫的抗性，还能通过调控宿主生理代谢与多样微生物群落的互作，塑造根际、叶际及种带微生物群落的结构与功能。本研究综述了香柱菌属内生真菌影响禾草根际（与土壤）、叶际和种带微生物群落组成与多样性的研究进展。系统介绍了香柱菌属内生真菌通过改善宿主水分利用效率、调节根系分泌物、影响叶片次生代谢产物及改变土壤理化性质等，选择性地促进或抑制宿主植物不同部位特定微生物类群的定殖，并普遍降低病原菌丰度和菌根侵染率的相关研究成果。进一步阐释了此类调控作用在代际间垂直传播，及其助力宿主植物适应胁迫环境的潜在机制。综上，本研究介绍了香柱菌属内生真菌在禾草微生物群落调控及生态功能中的关键作用，总结梳理了其在抗逆育种、微生物资源开发及可持续草地管理方面已经发挥或潜在的应用价值。

关键词: 禾草, 香柱菌属内生真菌, 根际, 叶际, 微生物

Abstract:

Epichlo? endophytes primarily colonize the aboveground tissues of host grasses， forming a stable， non-pathogenic symbiotic system. These endophytes not only enhance the host’s resistance to biotic and abiotic stresses but also shape the structure and function of rhizosphere， phyllosphere， and seed-associated microbial communities by regulating host physiological metabolism and interacting with diverse microbial populations. This review summarizes recent research on the effects of Epichlo? endophytes on the composition and diversity of microbial communities in the rhizosphere （and soil）， phyllosphere， and seeds of grasses. Present studies demonstrate that Epichlo? endophytes can selectively promote or inhibit the colonization of specific microbial groups in different plant compartments by improving host water-use efficiency， modulating root exudates， altering foliar secondary metabolites， and modifying soil physicochemical properties， while generally reducing pathogen abundance and mycorrhizal infection rates. This review further elaborates on the vertical transmission of these regulatory effects across plant generations and their potential mechanisms in enhancing host adaptation to environmental stresses. In conclusion， this review highlights the pivotal roles of Epichlo? endophytes in regulating grass-associated microbial communities and maintaining ecological functions， and summarizes their established and potential applications in stress-resilient breeding， microbial resource development， and sustainable grassland management.

Key words: grasses, Epichlo? endophyte, rhizosphere, phyllosphere, microorganisms

王美君, 崔龙轩, 南志标, 夏超. 香柱菌属内生真菌影响宿主禾草微生物群落结构与多样性的研究进展[J]. 草业学报, 2026, 35(10): 237-251.

Mei-jun WANG, Long-xuan CUI, Zhi-biao NAN, Chao XIA. Research progress on the effects of Epichloë endophytes on the structure and diversity of microbial communities in host grasses[J]. Acta Prataculturae Sinica, 2026, 35(10): 237-251.

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内生真菌 Endophytes	宿主 Host	作用 Function	参考文献 Reference
隐匿香柱菌 E. occultans	多花黑麦草 Lolium multiflorum	对根际土壤细菌的呼吸作用没有显著影响。Has no significant effect on the respiratory activity of rhizosphere soil bacteria.	［60］
		改变了宿主根际土壤细菌群落组成。Alters the composition of the host rhizosphere soil bacterial community.	［61］
		增加真菌活性，降低菌根真菌侵染率。Enhances fungal activity， and lower the mycorrhizal fungi infection rate.	［62］
黄化香柱菌 E. amarillans	美洲沙茅草 A. breviligulata	在水分胁迫条件下降低了植株根系相关细菌群落的多样性。Reduces the diversity of plant root-associated bacterial communities under water stress conditions.	［38］
		增加了根部内生真菌的丰富度并且在氮肥富集条件下改变了根部内生真菌群落的组成。Increases the abundance of root endophytic fungi and alters the composition of root endophytic fungal communities under nitrogen fertilizer enrichment conditions.	［63］
聚生香柱菌 E. coenophiala	高羊茅 F. arundinacea	增加了根际土壤细菌的多样性。Enhances the diversity of rhizosphere soil bacteria.	［41］
聚生香柱菌 E. coenophiala	高羊茅 F. arundinacea	降低了宿主在黏土生境下的革兰氏阳性细菌（G+）的多样性。Decreases the diversity of Gram’positive bacteria （G+） in the host under clay habitat conditions.	［64］
		对根部溶磷细菌多样性没有显著的相互作用。There is no significant interaction with the diversity of phosphate-solubilizing bacteria （PSB） in the roots.	［42］
		降低了古细菌的群落多样性，抑制了黏壤土中革兰氏阳性菌的生长。Reduces the community diversity of archaea and inhibits the growth of Gram-positive bacteria in clay loam.	［65］
	苇状羊茅Schedonorus arundinaceus	降低菌根真菌的定殖。Reduces the colonization of mycorrhizal fungi.	［66］
	苇状羊茅Schedonorus arundinaceus	在不同的土壤含水量下，对丛枝菌根真菌的定殖率不同。The colonization rate of AMF varies under different soil moisture contents.	［67］
		减少了根部的菌根定殖。Decreases mycorrhizal colonization in roots.	［29］
黑麦草香柱菌 E. festucae var. lolii	多年生黑麦草 L. perenne	显著增加了 16S rRNA 基因的绝对丰度以及土壤细菌群落的多样性。Significantly increases the absolute abundance of the 16S rRNA gene and the diversity of soil bacterial communities.	［68］
		改变了根际土壤中与氮循环相关的细菌基因丰度，尽管对土壤真菌群落的绝对丰度和多样性没有显著影响。Alters the abundance of bacterial genes related to nitrogen cycling in rhizosphere soil， although it has no significant effect on the absolute abundance and diversity of soil fungal communities.	［69］
甘肃香柱菌 E. gansuensis	醉马草 A. inebrians	减少了根际土壤中可培养细菌的种类数目。Decreases the number of culturable bacterial species in the rhizosphere soil.	［70］
		降低了根相关细菌多样性，增加了根际土壤细菌群落的多样性。Reduces root-associated bacterial diversity while increasing the diversity of rhizosphere soil bacterial communities.	［39］
		在盐胁迫下，显著增加了宿主养分提取和土壤微生物丰度。Significantly enhances host nutrient uptake and soil microbial abundance under salt stress.	［43］
		宿主抵抗CdCl?过程中，内生真菌增强了根际细菌物种间的协作。During the host’s resistance to CdCl?， strengthen the cooperation between rhizosphere bacterial species.	［71］
		在NH??-N处理下增加了宿主根部细菌的α-多样性。Increases the α-diversity of host root bacteria under NH?⁺-N treatment.	［72］
		降低了根系相关真菌群落的多样性，显著降低了根相关真菌多样性。Reduces the diversity of root-associated fungal communities， significantly reduces the diversity of root-associated fungi （RAF）.	［54］
		减少了根际土壤中真菌的数目。Decreases the number of fungi in rhizosphere soil.	［73］
		降低盐胁迫对真菌群落多样性的影响。Mitigates the impact of salt stress on fungal community diversity.	［74］
雀麦香柱菌 E. bromicola	羊草 Leymus chinensis	提高丛枝菌根真菌的侵染率和孢子密度。Improves the infection rate and spore density of arbuscular mycorrhizal fungi （AMF）.	［75］
		增加根际土壤中真菌的数目。Increases the number of fungi in rhizosphere soil.	［76］
雀麦香柱菌 E. bromicola	羊草 Leymus chinensis	增加根系和根际土壤中的真菌群落多样性和丰富度。Enhances the diversity and richness of fungal communities in roots and rhizosphere soil.	［77］
		随植物生长时间的延长，内生真菌与生长时间互作对真菌和细菌丰度影响显著。With the extension of plant growth period， the interaction between endophytic fungi and growth time has a significant impact on the abundance of fungi and bacteria.	［36］
		对宿主根部丛枝菌根真菌总定殖率没有影响。Has no effect on the total colonization rate of arbuscular mycorrhizal fungi （AMF） in host roots.	［53］
	高羊茅F. arundinacea	减少丛枝菌根真菌定殖率。Reduces the colonization rate of arbuscular mycorrhizal fungi （AMF）.	［56］
		降低菌根真菌的定殖。Reduces the colonization of mycorrhizal fungi.	［40］
		在盐碱胁迫条件下，显著增加了宿主根部植物促生真菌的多样性并改变了其组成。Significantly increases the diversity of plant growth-promoting fungi （PSE） in host roots and alters their composition under saline-alkali stress conditions.	［42］
	短芒大麦草Hordeum brevisubulatum	提高土壤中的氨氧化细菌丰度。Increases the abundance of ammonia-oxidizing bacteria in soil.	［78］
	布顿大麦草 Hordeum bogdanii	抑制了土壤真菌的丰富度，但提高了土壤真菌的多样性。Inhibits the abundance of soil fungi but improves the diversity of soil fungi.	［79］
		降低菌根真菌的定殖。Reduces the colonization of mycorrhizal fungi.	［80］
羊茅香柱菌 E. festucae	多年生黑麦草 L. perenne	显著增加了植物刈割返田后和植物自然生长土壤中真菌群落的多样性。 Significantly increases the diversity of fungal communities in soil after plant mowing and returning to the field， as well as in soil with natural plant growth.	［81］
	紫羊茅 Festuca rubra	显著提高根际土壤真菌群落的代谢途径和次生代谢产物生物合成功能。 Significantly improves the metabolic pathways and secondary metabolite biosynthesis functions of the rhizosphere soil fungal community.	［82］
潘佩香柱菌 E. panpeana	奥氏雀麦 Bromus auleticus	增加根际溶磷真菌多样性。Increases the diversity of phosphate-solubilizing fungi in the rhizosphere.	［59］
西伯利亚香柱菌 E. sibirica	西伯利亚羽茅Achnatherum sibiricum	降低菌根真菌的定殖。Reduces the colonization of mycorrhizal fungi.	［83］
滕布拉德香柱菌 E. tembladerae	刚毛雀麦 Bromus setifolius	增加菌根真菌的定殖。Increases the colonization of mycorrhizal fungi.	［58］
香柱菌 E. typhina	多年生黑麦草 L. perenne	降低菌根真菌的定殖。Reduces the colonization of mycorrhizal fungi.	［57］
香柱菌 E. typhina	多年生黑麦草 L. perenne	抑制菌根侵染率。Inhibits the mycorrhizal infection rate.	［84］
披碱草香柱菌 E. elymi	刺毛披碱草 Elymus hystrix	增加菌根真菌摩西球囊霉（Glomus mosseae）的定殖，降低菌根真菌明球囊霉（Glomus claroideum）的定殖。Increases the colonization of the mycorrhizal fungus G. mosseae and decreases the colonization of the mycorrhizal fungus G. claroideum.	［40］

内生真菌 Endophytes	宿主 Host	作用 Function	参考文献 Reference
隐匿香柱菌 E. occultans	多花黑麦草 Lolium multiflorum	对根际土壤细菌的呼吸作用没有显著影响。Has no significant effect on the respiratory activity of rhizosphere soil bacteria.	［60］
		改变了宿主根际土壤细菌群落组成。Alters the composition of the host rhizosphere soil bacterial community.	［61］
		增加真菌活性，降低菌根真菌侵染率。Enhances fungal activity， and lower the mycorrhizal fungi infection rate.	［62］
黄化香柱菌 E. amarillans	美洲沙茅草 A. breviligulata	在水分胁迫条件下降低了植株根系相关细菌群落的多样性。Reduces the diversity of plant root-associated bacterial communities under water stress conditions.	［38］
		增加了根部内生真菌的丰富度并且在氮肥富集条件下改变了根部内生真菌群落的组成。Increases the abundance of root endophytic fungi and alters the composition of root endophytic fungal communities under nitrogen fertilizer enrichment conditions.	［63］
聚生香柱菌 E. coenophiala	高羊茅 F. arundinacea	增加了根际土壤细菌的多样性。Enhances the diversity of rhizosphere soil bacteria.	［41］
聚生香柱菌 E. coenophiala	高羊茅 F. arundinacea	降低了宿主在黏土生境下的革兰氏阳性细菌（G+）的多样性。Decreases the diversity of Gram’positive bacteria （G+） in the host under clay habitat conditions.	［64］
		对根部溶磷细菌多样性没有显著的相互作用。There is no significant interaction with the diversity of phosphate-solubilizing bacteria （PSB） in the roots.	［42］
		降低了古细菌的群落多样性，抑制了黏壤土中革兰氏阳性菌的生长。Reduces the community diversity of archaea and inhibits the growth of Gram-positive bacteria in clay loam.	［65］
	苇状羊茅Schedonorus arundinaceus	降低菌根真菌的定殖。Reduces the colonization of mycorrhizal fungi.	［66］
	苇状羊茅Schedonorus arundinaceus	在不同的土壤含水量下，对丛枝菌根真菌的定殖率不同。The colonization rate of AMF varies under different soil moisture contents.	［67］
		减少了根部的菌根定殖。Decreases mycorrhizal colonization in roots.	［29］
黑麦草香柱菌 E. festucae var. lolii	多年生黑麦草 L. perenne	显著增加了 16S rRNA 基因的绝对丰度以及土壤细菌群落的多样性。Significantly increases the absolute abundance of the 16S rRNA gene and the diversity of soil bacterial communities.	［68］
		改变了根际土壤中与氮循环相关的细菌基因丰度，尽管对土壤真菌群落的绝对丰度和多样性没有显著影响。Alters the abundance of bacterial genes related to nitrogen cycling in rhizosphere soil， although it has no significant effect on the absolute abundance and diversity of soil fungal communities.	［69］
甘肃香柱菌 E. gansuensis	醉马草 A. inebrians	减少了根际土壤中可培养细菌的种类数目。Decreases the number of culturable bacterial species in the rhizosphere soil.	［70］
		降低了根相关细菌多样性，增加了根际土壤细菌群落的多样性。Reduces root-associated bacterial diversity while increasing the diversity of rhizosphere soil bacterial communities.	［39］
		在盐胁迫下，显著增加了宿主养分提取和土壤微生物丰度。Significantly enhances host nutrient uptake and soil microbial abundance under salt stress.	［43］
		宿主抵抗CdCl?过程中，内生真菌增强了根际细菌物种间的协作。During the host’s resistance to CdCl?， strengthen the cooperation between rhizosphere bacterial species.	［71］
		在NH??-N处理下增加了宿主根部细菌的α-多样性。Increases the α-diversity of host root bacteria under NH?⁺-N treatment.	［72］
		降低了根系相关真菌群落的多样性，显著降低了根相关真菌多样性。Reduces the diversity of root-associated fungal communities， significantly reduces the diversity of root-associated fungi （RAF）.	［54］
		减少了根际土壤中真菌的数目。Decreases the number of fungi in rhizosphere soil.	［73］
		降低盐胁迫对真菌群落多样性的影响。Mitigates the impact of salt stress on fungal community diversity.	［74］
雀麦香柱菌 E. bromicola	羊草 Leymus chinensis	提高丛枝菌根真菌的侵染率和孢子密度。Improves the infection rate and spore density of arbuscular mycorrhizal fungi （AMF）.	［75］
		增加根际土壤中真菌的数目。Increases the number of fungi in rhizosphere soil.	［76］
雀麦香柱菌 E. bromicola	羊草 Leymus chinensis	增加根系和根际土壤中的真菌群落多样性和丰富度。Enhances the diversity and richness of fungal communities in roots and rhizosphere soil.	［77］
		随植物生长时间的延长，内生真菌与生长时间互作对真菌和细菌丰度影响显著。With the extension of plant growth period， the interaction between endophytic fungi and growth time has a significant impact on the abundance of fungi and bacteria.	［36］
		对宿主根部丛枝菌根真菌总定殖率没有影响。Has no effect on the total colonization rate of arbuscular mycorrhizal fungi （AMF） in host roots.	［53］
	高羊茅F. arundinacea	减少丛枝菌根真菌定殖率。Reduces the colonization rate of arbuscular mycorrhizal fungi （AMF）.	［56］
		降低菌根真菌的定殖。Reduces the colonization of mycorrhizal fungi.	［40］
		在盐碱胁迫条件下，显著增加了宿主根部植物促生真菌的多样性并改变了其组成。Significantly increases the diversity of plant growth-promoting fungi （PSE） in host roots and alters their composition under saline-alkali stress conditions.	［42］
	短芒大麦草Hordeum brevisubulatum	提高土壤中的氨氧化细菌丰度。Increases the abundance of ammonia-oxidizing bacteria in soil.	［78］
	布顿大麦草 Hordeum bogdanii	抑制了土壤真菌的丰富度，但提高了土壤真菌的多样性。Inhibits the abundance of soil fungi but improves the diversity of soil fungi.	［79］
		降低菌根真菌的定殖。Reduces the colonization of mycorrhizal fungi.	［80］
羊茅香柱菌 E. festucae	多年生黑麦草 L. perenne	显著增加了植物刈割返田后和植物自然生长土壤中真菌群落的多样性。 Significantly increases the diversity of fungal communities in soil after plant mowing and returning to the field， as well as in soil with natural plant growth.	［81］
	紫羊茅 Festuca rubra	显著提高根际土壤真菌群落的代谢途径和次生代谢产物生物合成功能。 Significantly improves the metabolic pathways and secondary metabolite biosynthesis functions of the rhizosphere soil fungal community.	［82］
潘佩香柱菌 E. panpeana	奥氏雀麦 Bromus auleticus	增加根际溶磷真菌多样性。Increases the diversity of phosphate-solubilizing fungi in the rhizosphere.	［59］
西伯利亚香柱菌 E. sibirica	西伯利亚羽茅Achnatherum sibiricum	降低菌根真菌的定殖。Reduces the colonization of mycorrhizal fungi.	［83］
滕布拉德香柱菌 E. tembladerae	刚毛雀麦 Bromus setifolius	增加菌根真菌的定殖。Increases the colonization of mycorrhizal fungi.	［58］
香柱菌 E. typhina	多年生黑麦草 L. perenne	降低菌根真菌的定殖。Reduces the colonization of mycorrhizal fungi.	［57］
香柱菌 E. typhina	多年生黑麦草 L. perenne	抑制菌根侵染率。Inhibits the mycorrhizal infection rate.	［84］
披碱草香柱菌 E. elymi	刺毛披碱草 Elymus hystrix	增加菌根真菌摩西球囊霉（Glomus mosseae）的定殖，降低菌根真菌明球囊霉（Glomus claroideum）的定殖。Increases the colonization of the mycorrhizal fungus G. mosseae and decreases the colonization of the mycorrhizal fungus G. claroideum.	［40］

内生真菌 Endophytes	宿主 Host	作用 Function	参考文献 Reference
甘肃香柱菌 E. gansuensis	醉马草 A. inebrians	显著降低了种带微生物群落的丰富度和多样性。Significantly reduces the richness and diversity of the seed-borne microbial community.	［95］
		显著降低了种子内生和附生细菌群落的多样性。Significantly decreases the diversity of endophytic and epiphytic bacterial communities in seeds.	［96］
		改变不同世代之间细菌和真菌的群落组成。Alters the community composition of bacteria and fungi across different generations.	［102］
隐匿香柱菌 E. occultans	多花黑麦草 L. multiflorum	增加了细菌多样性，并使细菌群落更加均衡。Increases bacterial diversity and makes the bacterial community more balanced.	［93］
香柱菌 E. typhina	散穗雀麦 Puccinellia distans	影响真菌群落的组成。Affects the composition of the fungal community.	［105］
中华香柱菌 E. sinensis	中华羊茅 F.sinensis	对种带真菌多样性无显著影响。 Has no significant effect on the diversity of seed-borne fungi.	［106］

内生真菌 Endophytes	宿主 Host	作用 Function	参考文献 Reference
甘肃香柱菌 E. gansuensis	醉马草 A. inebrians	显著降低了种带微生物群落的丰富度和多样性。Significantly reduces the richness and diversity of the seed-borne microbial community.	［95］
		显著降低了种子内生和附生细菌群落的多样性。Significantly decreases the diversity of endophytic and epiphytic bacterial communities in seeds.	［96］
		改变不同世代之间细菌和真菌的群落组成。Alters the community composition of bacteria and fungi across different generations.	［102］
隐匿香柱菌 E. occultans	多花黑麦草 L. multiflorum	增加了细菌多样性，并使细菌群落更加均衡。Increases bacterial diversity and makes the bacterial community more balanced.	［93］
香柱菌 E. typhina	散穗雀麦 Puccinellia distans	影响真菌群落的组成。Affects the composition of the fungal community.	［105］
中华香柱菌 E. sinensis	中华羊茅 F.sinensis	对种带真菌多样性无显著影响。 Has no significant effect on the diversity of seed-borne fungi.	［106］