微生物学报

内生菌 Endophyte	VTE属 VTE genus	宿主植物 Plant	载体 Carrier	功能 Functions
内生细菌 Endophytic bacteria	泛菌属Pantoea	水稻Oryza sativa^[23] 玉米Zea mays^[24] 桉树Eucalyptus sp.^[16]	种子Seed 种子Seed 种子Seed	调节植物激素Regulate plant hormones^[38-40] 固氮作用Nitrogen fixation^[41] 产铁载体Produce siderophores^[38-39,41] 增强植物抗病性Enhance plant disease resistance^[42] 促进植物生长Promote plant growth^{[16,23,38-40,43]}
黄单胞菌Xanthomonas	水稻O. sativa^[23] 东南景天Sedum alfredii^[7]	种子Seed 扦插枝Cutting	固氮作用Nitrogen fixation^[23,44] 产铁载体Produce siderophores^[44] 促进植物生长Promote plant growth^[23]
肠杆菌属Enterobacter	水稻O. sativa^[40]	种子Seed	调节植物激素Regulate plant hormones^[40,45-46] 溶解磷能力Phosphorus solubilizing ability^[47] 促进植物生长Promote plant growth^[24,40]
假单胞菌属Pseudomonas	水稻O. sativa^[48]	种子Seed	调节植物激素Regulate plant hormones^[49] 固氮作用Nitrogen fixation^[50-52] 产铁载体Produce siderophores^{[47,49,53-54]} 溶解磷能力Phosphorus solubilizing ability^[55-58] 增强植物抗病性Enhance plant disease resistance^[59-60]
鞘氨醇单胞菌属Sphingomonas	水稻O. sativa^[48] 葡萄Vitis vinifera^[8]	种子Seed 茎尖Shoots	-
新鞘氨醇杆菌属Novosphingobium	葡萄V. vinifera^[8]	茎尖Shoots	-
微杆菌属Microbacterium	水稻O. sativa^[20] 柳枝稷Panicum virgatum^[17]	种子Seed 种子Seed	-
类芽孢杆菌属Paenibacillus	桉树Eucalyptus sp.^[16] 玉米Z. mays^[24]	种子Seed 种子Seed	促进植物生长Promote plant growth^[16,61] 增强植物抗病性Enhance plant disease resistance^[26,62-63]
腐殖质类芽孢杆菌Paenibacillus humicus	桉树Eucalyptus sp.^[16]	种子Seed	-
蒙氏肠球菌Enterococcus mundtii	桉树Eucalyptus sp.^[16]	种子Seed	-
甲基杆菌属Methylobacterium	桉树Eucalyptus sp.^[16]	种子Seed	促进植物生长Promote plant growth^[64-67] 提高重金属耐受Enhance heavy metal tolerance^[67]
梭菌属Clostridium	玉米Z. mays^[24]	种子Seed	-
中生根瘤菌属Mesorhizobium	白刺花Sophora davidii^[68]	种子Seed	增强植物抗逆性Enhance plant stress resistance^[68]
贪铜菌属Cupriavidus	葡萄V. vinifera^[8]	茎尖Shoots	-
劳尔氏菌属Ralstonia	葡萄V. vinifera^[8]	茎尖Shoots	-
芽孢杆菌属Bacillus	水稻O. sativa^[6] 桉树Eucalyptus sp.^[16] 柳枝稷P. virgatum^[17] 仙人掌Opuntia stricta^[50] 番茄Solanum lycopersicum^[69] 酢浆草Oxalis corniculate^[70] 葡萄V. vinifera^[1,8,12]	种子Seed 种子Seed 种子Seed 种子Seed 种子Seed 种子Seed 种子Seed, 扦插枝Cutting	调节植物激素Regulate plant hormones^[71-74] 固氮作用Nitrogen fixation^[50,72] 产铁载体Produce siderophores^[56,75] 溶解磷能力^[53,56-57] 增强植物抗病性Enhance plant disease resistance^{[39,69,76-79]} 促进植物生长Promote plant growth^[39,72,80]
内生真菌 Endophytic fungi	链格孢属Alternaria	欧洲千里光Senecio vulgaris^[34] 长叶车前Plantago lanceolata^[34] 酸模Rumex acetosa^[34] 紫茎泽兰Ageratina adenophora^[81] 葡萄V. vinifera^[8]	种子Seed 种子Seed 种子Seed 种子Seed 茎尖Shoots	促进植物生长Promote plant growth^[63]
Epichloё (Neotyphodium)	禾本科Poaceae^[82] 加拿大黑麦Elymus canadensis^[83] 醉马草Achnatherum inebrian^[35] 野大麦Hordeum brevisubulatum^[35] 高羊茅Festuca arundinacea^[35] 黑麦草Lolium multiflorum^[36] Poa lanuginosa poir^[33] Poe bonariensis^[33]	种子Seed 种子Seed 种子Seed 种子Seed 种子Seed 种子Seed 种子Seed 种子Seed	促进植物生长Promote plant growth^[41,84-85] 改善植物光合作用Improve plant photosynthesis^[41,84] 增强植物抗逆性Enhance plant stress resistance^[86-87] 生产麦角碱和喹啉生物碱Produce ergot alkaloids and quinoline alkaloids^[88]
白僵菌属Beauveria	辐射松Pinus radiata^[89]	种子Seed	-
白粉菌属Erysiphe	葡萄V. vinifera^[8]	茎尖Shoots	-
根瘤菌Rhizosphaera	东南景天S. alfredii^[7] 葡萄V. vinifera^[8]	扦插枝Cutting 茎尖Shoots	固氮作用Nitrogen fixation^[39,90-91] 溶解磷能力Phosphorus solubilizing ability^[47,92] 增强植物抗逆性Enhance plant stress resistance^[92-93]
Davidiella	葡萄V. vinifera^[8]	茎尖Shoots	-
隐球菌属Cryptococcu	紫茎泽兰A. adenophora^[81]	种子Seed	-
枝孢属Cladosporiu	矢车菊Centaurea Cyanus^[34] 罂粟Papaver rhoeas^[34] 黑矢车菊Centaurea nigra^[34] 紫茎泽兰A. adenophora^[81] 葡萄V. vinifera^[8]	种子Seed 种子Seed 种子Seed 种子Seed 茎尖Shoots	促进植物生长Promote plant growth^[94] 抗氧化能力Antioxidant capacity^[95]
内生放线菌 Endophytic actinomycetes	链霉菌属Streptomyces	东南景天S. alfredii^[7]	扦插枝Cutting	促进植物生长Promote plant growth^[7] 增强植物抗逆性Enhance plant stress resistance^[7]
类诺卡氏属Nocardiopsis	东南景天S. alfredii^[7]	扦插枝Cutting	-
假诺卡氏菌属Pseudonocardia	东南景天S. alfredii^[7]	扦插枝Cutting	-

内生菌 Endophyte	VTE属 VTE genus	宿主植物 Plant	载体 Carrier	功能 Functions
内生细菌 Endophytic bacteria	泛菌属Pantoea	水稻Oryza sativa^[23] 玉米Zea mays^[24] 桉树Eucalyptus sp.^[16]	种子Seed 种子Seed 种子Seed	调节植物激素Regulate plant hormones^[38-40] 固氮作用Nitrogen fixation^[41] 产铁载体Produce siderophores^[38-39,41] 增强植物抗病性Enhance plant disease resistance^[42] 促进植物生长Promote plant growth^{[16,23,38-40,43]}
黄单胞菌Xanthomonas	水稻O. sativa^[23] 东南景天Sedum alfredii^[7]	种子Seed 扦插枝Cutting	固氮作用Nitrogen fixation^[23,44] 产铁载体Produce siderophores^[44] 促进植物生长Promote plant growth^[23]
肠杆菌属Enterobacter	水稻O. sativa^[40]	种子Seed	调节植物激素Regulate plant hormones^[40,45-46] 溶解磷能力Phosphorus solubilizing ability^[47] 促进植物生长Promote plant growth^[24,40]
假单胞菌属Pseudomonas	水稻O. sativa^[48]	种子Seed	调节植物激素Regulate plant hormones^[49] 固氮作用Nitrogen fixation^[50-52] 产铁载体Produce siderophores^{[47,49,53-54]} 溶解磷能力Phosphorus solubilizing ability^[55-58] 增强植物抗病性Enhance plant disease resistance^[59-60]
鞘氨醇单胞菌属Sphingomonas	水稻O. sativa^[48] 葡萄Vitis vinifera^[8]	种子Seed 茎尖Shoots	-
新鞘氨醇杆菌属Novosphingobium	葡萄V. vinifera^[8]	茎尖Shoots	-
微杆菌属Microbacterium	水稻O. sativa^[20] 柳枝稷Panicum virgatum^[17]	种子Seed 种子Seed	-
类芽孢杆菌属Paenibacillus	桉树Eucalyptus sp.^[16] 玉米Z. mays^[24]	种子Seed 种子Seed	促进植物生长Promote plant growth^[16,61] 增强植物抗病性Enhance plant disease resistance^[26,62-63]
腐殖质类芽孢杆菌Paenibacillus humicus	桉树Eucalyptus sp.^[16]	种子Seed	-
蒙氏肠球菌Enterococcus mundtii	桉树Eucalyptus sp.^[16]	种子Seed	-
甲基杆菌属Methylobacterium	桉树Eucalyptus sp.^[16]	种子Seed	促进植物生长Promote plant growth^[64-67] 提高重金属耐受Enhance heavy metal tolerance^[67]
梭菌属Clostridium	玉米Z. mays^[24]	种子Seed	-
中生根瘤菌属Mesorhizobium	白刺花Sophora davidii^[68]	种子Seed	增强植物抗逆性Enhance plant stress resistance^[68]
贪铜菌属Cupriavidus	葡萄V. vinifera^[8]	茎尖Shoots	-
劳尔氏菌属Ralstonia	葡萄V. vinifera^[8]	茎尖Shoots	-
芽孢杆菌属Bacillus	水稻O. sativa^[6] 桉树Eucalyptus sp.^[16] 柳枝稷P. virgatum^[17] 仙人掌Opuntia stricta^[50] 番茄Solanum lycopersicum^[69] 酢浆草Oxalis corniculate^[70] 葡萄V. vinifera^[1,8,12]	种子Seed 种子Seed 种子Seed 种子Seed 种子Seed 种子Seed 种子Seed, 扦插枝Cutting	调节植物激素Regulate plant hormones^[71-74] 固氮作用Nitrogen fixation^[50,72] 产铁载体Produce siderophores^[56,75] 溶解磷能力^[53,56-57] 增强植物抗病性Enhance plant disease resistance^{[39,69,76-79]} 促进植物生长Promote plant growth^[39,72,80]
内生真菌 Endophytic fungi	链格孢属Alternaria	欧洲千里光Senecio vulgaris^[34] 长叶车前Plantago lanceolata^[34] 酸模Rumex acetosa^[34] 紫茎泽兰Ageratina adenophora^[81] 葡萄V. vinifera^[8]	种子Seed 种子Seed 种子Seed 种子Seed 茎尖Shoots	促进植物生长Promote plant growth^[63]
Epichloё (Neotyphodium)	禾本科Poaceae^[82] 加拿大黑麦Elymus canadensis^[83] 醉马草Achnatherum inebrian^[35] 野大麦Hordeum brevisubulatum^[35] 高羊茅Festuca arundinacea^[35] 黑麦草Lolium multiflorum^[36] Poa lanuginosa poir^[33] Poe bonariensis^[33]	种子Seed 种子Seed 种子Seed 种子Seed 种子Seed 种子Seed 种子Seed 种子Seed	促进植物生长Promote plant growth^[41,84-85] 改善植物光合作用Improve plant photosynthesis^[41,84] 增强植物抗逆性Enhance plant stress resistance^[86-87] 生产麦角碱和喹啉生物碱Produce ergot alkaloids and quinoline alkaloids^[88]
白僵菌属Beauveria	辐射松Pinus radiata^[89]	种子Seed	-
白粉菌属Erysiphe	葡萄V. vinifera^[8]	茎尖Shoots	-
根瘤菌Rhizosphaera	东南景天S. alfredii^[7] 葡萄V. vinifera^[8]	扦插枝Cutting 茎尖Shoots	固氮作用Nitrogen fixation^[39,90-91] 溶解磷能力Phosphorus solubilizing ability^[47,92] 增强植物抗逆性Enhance plant stress resistance^[92-93]
Davidiella	葡萄V. vinifera^[8]	茎尖Shoots	-
隐球菌属Cryptococcu	紫茎泽兰A. adenophora^[81]	种子Seed	-
枝孢属Cladosporiu	矢车菊Centaurea Cyanus^[34] 罂粟Papaver rhoeas^[34] 黑矢车菊Centaurea nigra^[34] 紫茎泽兰A. adenophora^[81] 葡萄V. vinifera^[8]	种子Seed 种子Seed 种子Seed 种子Seed 茎尖Shoots	促进植物生长Promote plant growth^[94] 抗氧化能力Antioxidant capacity^[95]
内生放线菌 Endophytic actinomycetes	链霉菌属Streptomyces	东南景天S. alfredii^[7]	扦插枝Cutting	促进植物生长Promote plant growth^[7] 增强植物抗逆性Enhance plant stress resistance^[7]
类诺卡氏属Nocardiopsis	东南景天S. alfredii^[7]	扦插枝Cutting	-
假诺卡氏菌属Pseudonocardia	东南景天S. alfredii^[7]	扦插枝Cutting	-

植物垂直传递内生菌研究进展

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郭丽榕 ¹ , 郭青钰 ¹ , 朱书生 ² , 杨明挚 ¹^,^*

微生物学报 | 综述 2025,65(2): 489-504

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微生物学报 | 综述 2025, 65(2): 489-504

植物垂直传递内生菌研究进展

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郭丽榕¹, 郭青钰¹, 朱书生², 杨明挚¹^,^*

作者信息

1 云南大学生态与环境学院，云南高原山地生态与退化环境修复重点实验室，云南昆明

2 云南农业大学植物保护学院，云南昆明

Research advances in vertically transmitted endophytes in plants

Lirong GUO¹, Qingyu GUO¹, Shusheng ZHU², Mingzhi YANG¹^,^*

Affiliations

1 Yunnan Key Laboratory for Plateau Mountain Ecology and Restoration of Degraded Environments, School of Ecology and Environmental Science, Yunnan University, Kunming, Yunnan, China

2 College of Plant Protection, Yunnan Agricultural University, Kunming, Yunnan, China

出版时间: 2025-02-04 doi: 10.13343/j.cnki.wsxb.20240527

文章导航

摘要

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植物内生菌因其与宿主间在空间和功能上的特殊关系，被称为植物的第二基因组，也是植物学和微生物学研究中的共同热点。植物中的内生菌依据其来源方式可分为水平传递内生菌(horizontally transmitted endophytes, HTE)和垂直传递内生菌(vertically transmitted endophytes, VTE)。其中，垂直传递内生菌有机会通过繁殖体(包括有性繁殖体和无性繁殖体)进一步传递到后代植株产生相应的宿主效应。VTE在宿主植物中的行为和功能更类似于宿主的一种获得性 “可遗传”性状，可应用于改良植物性状的生产实践中。因此，针对植物垂直传递内生菌的深入了解和研究无疑具有重要的理论和现实意义。本文综述了目前有关植物VTE的类群、多样性、传递及生理生态功能等方面的研究现状，并对VTE的研究和应用前景进行了展望。

关键词

垂直传递内生菌 / 多样性 / 种子内生菌 / 宿主效应 / 无性繁殖体

Abstract

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Due to the unique spatial and functional relationship with the host, plant endophytes are known as the second genome of plants and have become a research hotspot of plant sciences and microbiology. According to the origins, plant endophytes can be classified into horizontally transmitted endophytes (HTE) and vertically transmitted endophytes (VTE). Among them, VTEs have the potential to be further transmitted into later-generation plants through propagules (both sexual and asexual propagules) to produce host effects. The behaviors and functions of VTE along host generations are likely to be an acquired “heritable” trait of the host that can be applied in agricultural practices for plant trait improvement. Therefore, in-depth understanding and research on plant VTEs are of great theoretical and practical importance. We reviewed the research advances in the diversity, transmission, and physiological and ecological functions of plant VTEs and provide perspectives for future research and application of VTE resources.

Key words

vertically transmitted endophytes (VTE) / diversity / seed endophytes / host effects / asexual propagules

引用本文

郭丽榕, 郭青钰, 朱书生, 杨明挚. 植物垂直传递内生菌研究进展. 微生物学报, 2025 , 65 (2) : 489 -504 . DOI: 10.13343/j.cnki.wsxb.20240527

Lirong GUO, Qingyu GUO, Shusheng ZHU, Mingzhi YANG. Research advances in vertically transmitted endophytes in plants[J]. Acta Microbiologica Sinica, 2025 , 65 (2) : 489 -504 . DOI: 10.13343/j.cnki.wsxb.20240527

正文

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植物内生菌(endophyte)是指那些在其生活史的全部或部分阶段栖息于植物组织和器官内部的所有微生物的统称，而不考虑其功能^[1]。可培养内生菌(culturable endophytes)可以从经过表面消毒的植物器官(如根、茎、叶、花、果实和种子)中分离获得，而绝大多数的植物内生菌属于难培养内生菌(cultivation-recalcitrant endophytes)，通过分子生物学技术可以检测到这些内生菌的存在^[2]。

植物内生菌能够通过水平传递的方式从环境中获得，即：水平传递内生菌(horizontally transmitted endophytes, HTE)；另一种则是通过垂直传递的方式从母代植物中获得，即：垂直传递内生菌(vertically transmitted endophytes, VTE)^[1,3]。依据内生菌与宿主植物之间的关系，可以进一步将它们划分为专性内生菌(obligate endophytes)和兼性内生菌(facultative endophytes)。专性内生菌对宿主植物具有较高的依赖性，也是垂直传递内生菌的重要组成部分，这些微生物的整个生命周期都生活在宿主内，难以在其他环境中独立生存^[4]。当专性内生菌与宿主植物间建立起稳定且互惠互利的关系之后，这些内生菌就会在该植物的世代之间垂直传播，以确保这种互利关系的持续性，并在种子萌发及植物成长的各个阶段中发挥重要生物学功能^[5]。

内生菌的垂直传递可以通过种子进行^[6]，同时，它们也能借助无性繁殖体，如扦插枝条、块茎、鳞茎等^[7-9]作为媒介。种子是植物繁殖以及农业生产中的核心要素^[10]。大量的研究表明，植物种子内含有多样且丰富的内生微生物，即种子内生菌(seed endophytes, SE)。基于多组学的分析揭示，植物种子含有的有益且与植物基因型相关的特异性内生菌可以垂直传播至下一代^[11]。此外，以无性繁殖方式为主的作物，如葡萄、红薯、马铃薯等^[12]，其无性繁殖体中的内生菌也能够以垂直传递的方式传递给其子代无性繁殖的植株。同时，种子以及无性繁殖体中的内生菌可以通过子代植物的根系进入根际，参与构建根际土壤微生物群落，形成有益的植物相关微生物群落(plant association microbiome)，从而增强植物的生长和对环境的适应性^[7]。同样，源于种子或无性繁殖体的内生菌也能转移到植物体表，参与组装植物叶际微生物群落(phyllosphere microflora)^[13-14]。因此，VTE在构建子代植物相关微生物群落以及帮助宿主植物适应环境和开拓新的生态位中扮演着重要角色。

1 种子介导的VTE

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1.1 种子介导的细菌VTE

种子是内生菌在宿主植物间垂直传递的主要媒介，植物内生细菌的垂直传播机制已通过种子与幼苗内生菌群的重叠关系得到了验证^[15-16]，Gagne-Bourgue等^[17]在柳枝稷(Panicum virgatu)种子中发现芽孢杆菌(Bacillus)和微杆菌(Microbacterium)，这些内生细菌不仅存在于种子中，还能在由这些种子发育而来的子代植株体内持续存在并被检测到。Cope-Selby等^[18]在研究种子内生菌的来源时发现，芒草(Miscanthus sinensis)种子中的内生细菌主要来源于其亲本内生细菌。随着植物的生长，根际以及土壤中的少量细菌也可以侵入植物成为内生菌，进而转移到芒草种子内部，成为种子内生菌。Thomas等^[19]也观察到了番茄内生细菌的垂直传播现象。

水稻(Oryza sativa)是研究种子内生菌垂直传递较多的作物之一。Walitang等^[20]通过Sörensen指数和非度量多维尺度分析(nonmetric multidimensional scaling, NMDS)发现，子代种子中所携带的内生细菌与其亲代种子之间存在高度的相似度。这意味着子代种子内生细菌的来源具有多元性，各部分均有所贡献，但亲代种子仍是其最主要的来源；Hardoim等^[6]发现，从子代种子中分离出的嗜麦芽窄养单胞菌(Stenotrophomonas maltophilia)和嗜冷杆菌属(Ochrobactrum)细菌均来自上一代种子；水稻种子内生菌是子代植株内生细菌的重要来源，它们通过水稻种子传递给下一代，并成为成熟植物体内的优势内生菌物种^[21]。Wang等^[22]采用Illumina高通量测序技术调查了水稻芽、根、茎的内生细菌和真菌群落，发现水稻芽内共生的假单胞菌属(Pseudomonas)和鞘氨醇单胞菌属(Sphingomonas)细菌在茎和根内也被检测到。大量研究均表明，亲代水稻种子中的内生菌能够伴随植株的生长发育过程传播到根和茎，并最终进入子代种子，即实现了内生菌从亲代种子到子代种子的垂直传播^[23]。

针对玉米(Zea mays)种子内生菌的研究也得出了类似的结果。利用绿色荧光蛋白(green fluorescent protein, GFP)标记内生菌的实验发现，泛菌属(Pantoea)和假单胞菌属从玉米种子进入根和根际，进而转移到植物的其他组织，并且2个直系世代之间内生菌的垂直传播率高达44%^[24]，这表明内生菌可能在整个植物体内进行系统性传递。Liu等^[25]发现，玉米种子的内生细菌种类从亲本到子代具有连续性。通过分析连续2年内收获的玉米种子，均发现了泛菌属菌株的存在，这显示了种子垂直传递内生菌的相对稳定性^[26]。同样地，Ferreira等^[16]通过向桉树(Eucalyptus)种子接种经过绿色荧光蛋白(GFP)标记的成团泛菌(Pantoea agglomerans)，在种子发芽成长为幼苗的过程中，成功地在幼苗根部细胞间隙以及茎部木质部导管内追踪到了这些被标记的细菌。这一发现为内生菌能够从种子阶段垂直传递到幼苗阶段提供了直接的证据。

1.2 种子介导的真菌VTE

目前认为疯草内生真菌主要依靠种子带菌的方式传播到下一代^[27]。Neotyphodium属内生真菌被普遍认为是Epichloë属的无性世代^[28]。Neotyphodium属真菌能够在植物的叶片、花序和种子内共生，从而有机会实现垂直传播到子代植株^[29]。Epichloëcoenophiala、Epichloëuncinata及其杂交种通常以垂直传递的方式，通过植物种子进行无性传播^[30]。Tadych等^[31]发现，侵染偏生早熟禾亚种(Poa secunda subsp. Juncifolia)的Epichloë typhina subsp. poae不仅可以稳定地垂直传播，还可以通过产孢进行水平传播。中华羊茅(Festuca sinensis)内生真菌的传播方式也主要依赖种子的垂直传播^[32]。内生真菌Epichloё能在加拿大野黑麦(Elymus canadensis)中进行垂直传播，并在此过程中产生稳定的遗传变异^[33]。

此外，Hodgson等^[34]的研究发现，6种草本植物均能通过种子向后代植物实现垂直传递内生真菌；进一步的研究还发现，内生真菌如链格孢菌(Alternaria alternata)、球形枝孢菌(Cladosporium sphaerospermum)等，还在这些草本植物的花粉粒表面及其内部被检测到，这表明内生真菌能够在花粉管中定居并存活，极有可能成为内生菌进入正在发育种子的潜在途径^[34]。这一系列的发现有力地证实了种子内生菌的垂直传递确实是一种广泛存在的机制。

依据目前的研究数据，虽然内生菌在植物间的跨代传递确为一种普遍的现象，但子代植物及其种子中的绝大多数内生菌并非源自垂直传递方式^[17,24]，而是来源于水平传递的微生物。显然，通过水平传播方式获得的部分种子内生菌将有可能成为该宿主新的VTE类群。因此，植物中VTE类群的动态变化体现了宿主-内生菌间的相互选择过程(图1)，是植物-微生物相互作用中的核心环节，其机制值得进一步深入研究。

2 无性繁殖体介导的VTE

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理论上，由无性繁殖体介导的垂直传递内生菌类似于内生菌在宿主体内的系统性传递方式。无性繁殖体，如扦插枝，往往是子代植株的一部分，它们与新生的植物组织器官以及未来的繁殖体(枝条)之间缺乏明显的隔离，因此更易于实现垂直传递。Vannier等^[9]以一种草本植物(Glechoma hederacea)为模型，验证了内生菌在无性克隆后代中的垂直传递，并深入探讨了在克隆植物中共享内生菌的群体遗传特性：在严格控制环境微生物的条件下，母代植株中的大部分内生细菌和真菌都会垂直传播给由母代植株根萌发形成的子代植株，而古生菌则并未传递给后代；该研究证实，在内生菌通过无性繁殖进行垂直传播的过程中，存在一种宿主过滤机制。Luo等^[7]利用东南景天扦插枝建立了垂直传递内生菌-植株内共生体系，发现通过东南景天扦插枝垂直传播而建立的内生菌群至少持续了4个月；内生菌群可以从茎部传播到根内层，然后在新形成的根际中增殖；新产生的芽内生菌由自然适应宿主的内生菌类群组成，在植物发育过程中达到相对稳定的状态，并持续存在。

Xiang等^[8]利用高通量测序技术，追踪研究了酿酒葡萄品种玫瑰蜜(Vitis vinifera-V. labrusca)和赤霞珠(Vitis vinifera L.)从茎尖诱导的无菌组培苗，经炼苗、移栽至成熟植株不同阶段的内生菌群动态变化，发现随着葡萄的生长，组培苗中的核心内生菌能够迁移并定殖于克隆植株的根部、茎部和叶片中，结果表明，内生菌由母株葡萄藤茎尖垂直传递到由该茎尖诱导的组培苗及其不断继代的离体培养植株(in vitro-cultured plantlets, IVCP)，类似于种子传播的VTEs。Chen等^[12]将从葡萄植株中分离培养的VTE菌株ZX-2 (Bacillus cereus)成功导入到玫瑰蜜品种的葡萄扦插枝，并将这些扦插枝培养成葡萄植株，发现导入的ZX-2菌株对宿主葡萄植株内生菌群结构产生了显著影响，并在宿主葡萄植株不同部位迁徙，也出现在第二年萌发的新梢中；同时，该研究也证明扦插枝前期定殖的其他内生菌也可以传递给后代，这为有目的地改良葡萄扦插枝的VTE菌群，进而优化后代克隆群体的内生菌群结构提供了重要依据。

3 植物VTE的多样性

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3.1 植物VTE的分离鉴定

目前，针对植物VTE的鉴定，主要依据是该内生菌的物种、菌株或扩增子序列(amplicon sequence variant, ASV)是否同时在该植物的繁殖体和子代植株中存在。通过荧光标记来鉴定VTE^[8,24]是一种相对直接的方式，但这种方法一般仅局限于可培养的内生菌，对VTE和非VTE在分类学和生物学特征上的差异研究仍较为匮乏。从表型上看，内生菌必须具备在植株体内系统传递的能力，才有可能进入植物种子或其他繁殖体，并进一步从繁殖体迁移到子代植株的不同器官和组织中，实现垂直传递。因此，含有移动元件(contain mobile elements)应该是VTE的一个非常重要的前提条件。值得一提的是，从某一宿主中分离并鉴定的VTE菌株，在另一宿主中是否依然保持垂直传递的特性，还需要进一步的验证。

3.2 植物VTE的物种多样性

植物垂直传递内生菌种类繁多，大类群上与内生菌的多样性相一致，包括垂直传递的细菌、真菌以及放线菌。

目前，从东南景天(Sedum alfredii)^[7]、桉树(Eucalyptus sp.)^[16]、葡萄(Vitis vinifera)^[8]、水稻(Oryza sativa)^[23]、玉米(Zea mays)^[24]、柳枝稷(Panicum virgatum)^[17]等植物的种子及无性繁殖体中分离并鉴定出了多种垂直传递内生菌。同时，研究人员在醉马草(Achnatherum inebrians)^[35]、黑麦草(Lolium multiflorum)^[36]、野大麦(Hordeum brevisubulatum)^[35]、鹅观草(Roegneria kamoji)^[37]等禾本科植物中，则分离得到了可垂直传递的禾草Epichloë 内生真菌。

在属水平上，植物垂直传递内生细菌主要集中于假单胞菌属(Pseudomonas)、泛菌属(Pantoea)、肠杆菌属(Enterobacter)、微杆菌属(Microbacterium)、类芽孢杆菌属(Paenibacillus)、芽孢杆菌属(Bacillus)等，而植物垂直传递的内生真菌则主要有链格孢属(Alternaria)、枝孢属(Cladosporium)、球腔菌属(Mycosphaerella)、隐球菌属(Cryptococcus)以及Epichloë属等。相较于内生细菌和内生真菌，关于植物垂直传递内生放线菌的研究较为匮乏。科研人员目前在东南景天中成功分离并鉴定出了几种能够实现垂直传递的内生放线菌，包括链霉菌属(Streptomyces)、类诺卡氏属(Nocardioides)、假诺卡氏菌属(Pseudonocardia)等(表1)。

3.3 植物VTE的功能多样性

VTE属于植物内生菌中的特殊类群，该类群内生菌除了物种上的多样性外，功能上也具有多样性(表1)。

植物内生菌在促进宿主植物生长方面扮演着重要角色。首先，内生菌能够增强宿主植物对关键营养元素如氮、磷和铁的吸收能力，从而为植物提供必要的养分支持。其次，内生菌还能调节植物激素水平，这是其促进植物生长的另一主要途径。内生菌具备合成一种或多种关键植物激素的能力，包括生长素、细胞分裂素和赤霉素等，这些激素在调节植物生长和发育过程中发挥着至关重要的作用。另外，一些植物内生菌通过产生1-氨基环丙烷-1-羧酸脱氨酶(1-aminoeyclopropane-1-earboxylate-deaminase，ACC脱氨酶，简称ACCD酶)来抑制植物的乙烯水平，进而促进植物生长^[96]。泛菌属内生菌通过溶磷，产吲哚乙酸(indole-3-acetic acid, IAA)、铁载体、氢氰酸(hydrocyanic acid, HCN)等方式促进植物生长，是典型的植物促生长细菌^[38]。假单胞菌和芽孢杆菌是诱导宿主植物系统抗性的微生物类群，同时，这2个菌属的细菌也是常见的植物垂直传递内生菌重要来源^[1]，许多研究表明，这2个菌群在增强植物抗病性^{[39,59-60,76-77]}与促进植物生长^[39,72,80]方面均有显著效果。此外，从水稻种子中分离出的类芽孢杆菌、泛菌、微杆菌、假单胞菌等属的内生细菌具有抗真菌活性^[97]。对植物内生菌VTE的研究发现，大部分对植物产生有利效应的内生细菌都具有垂直传递特性，这反映了效应驱动的宿主-VTE选择机制。

禾草内生真菌能够影响植物病原真菌的生长，进而控制植物病害的发生与传播。内生真菌Epichloë能够产生具有抗虫活性的麦角类、吲哚二萜类、黑麦草碱和波胺这4种生物碱，这些生物碱对哺乳动物同样具有神经调节活性^[35]。此外，内生真菌Epichloë (Neotyphodium)或其与其寄主形成的共生体对多种病原真菌的生长具有显著的抑制作用^[98]。因此，垂直传递的Epichloë属内生真菌不仅能够提升宿主植物的抗逆性，还能促进宿主禾草的生长，从而增强宿主在自然植物群落中的竞争力。

放线菌是产生抗生素和其他生物活性物质最多的微生物类群^[99]。从喜树种子中分离的链霉菌属(Streptomyces)内生放线菌，能够抑制立枯丝核菌(Rhizoctonia solani Kühn)菌核的形成，并同时抑制其他一些对宿主有害的细菌类群的增殖^[100]。尽管该研究未明确这些种子内生菌是否具备垂直传递的能力，但在东南景天中发现的链霉菌属内生菌却具有垂直传递的特性^[7]。

垂直传递的内生菌不仅可以通过调节植物激素、产生铁载体等方式来促进植物生长，还能够改变宿主应对生物和非生物胁迫的能力(表1)。这进一步拓宽了垂直传递内生菌在农业生产中的利用价值。

4 VTE来源及其在子代植株的分布

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子代种子的大部分内生细菌都可以追溯到其亲本来源。赵霞^[101]研究了子代水稻种子内生细菌的来源，发现种子内生细菌中来源于亲代种子、根际、根围土、根和茎的微生物的比例分别为8.6%、0.3%、4.7%、8.5%和25.2%，还有12.8%的种子内生菌来源未知。该研究明确了种子内生细菌主要源自其亲代种子。Hardoim等^[6]指出，水稻种子内生细菌会首先在水稻根际定殖，然后转移到芽中。在整个植物发育过程中，茎部组织的内生细菌丰富度高于根组织。Wang等^[22]则发现，在健康植物组织中，真菌和细菌微生物群可以从芽传播到茎和根。与芽中的内生微生物群相比，根中的细菌丰富度增加，茎中的细菌丰富度减少，根系和茎部真菌丰富度随植株发育而减少。接种菌株PsJN (Burkholderia sp.)后的葡萄植株，在后续葡萄植株的根表面、花序、种子等部位均能检测到该菌株的存在，这表明了微生物从根系进入植物体后向植物地上部分的传递途径^[102]。栖息于宿主细胞壁中的Epichloë属内生真菌，其菌丝体会伴随宿主植物的生长逐渐延伸至植物叶鞘、叶片，甚至进入花序和种子等植物的地上部分，然后这些内生真菌会随着种子的萌发继续传播，从而感染新的植物^[103]。Chen等^[12]测定了蜡样芽孢杆菌ZX-2侵染的葡萄扦插枝的不同部位，发现蜡样芽孢杆菌ZX-2对葡萄扦插枝的侵染首先发生在木质部，从形态学下端向形态学上端传递，随后水平扩散至髓部和韧皮部。Xiang等^[8]通过研究2个不同葡萄品种(赤霞珠和玫瑰蜜)组培苗中内生细菌和真菌ASV在不同发育阶段的动态演变，发现在2个品种的4个生长阶段中，内生细菌的丰度和ASV数量均在第2阶段达到峰值，然后在第3阶段和第4阶段呈现下降趋势。在第4阶段，2个品种的根部ASV数量最多，且数量沿根、茎和叶方向呈逐渐递减。此外，该研究还发现，源于组培苗前期定殖的核心ASV (c-ASVs)在后期葡萄植株中呈现组织特异性分布模式(图2)。在内生细菌c-ASVs中，Bc01/Br02 (Cupriavidus)、Bc02/Br03 (Ralstonia)和Bc03/Br04 (Sphingomonas)在叶片和茎段中占主导地位，而Bc32/Br48 (Novosphingobium)在根段中占优势(图2A)。内生真菌c-ASVs Fc20 (Erysiphe)、Fc16 (Cladosporium)、Fc10 (Davidiella)、Fc48 (Alternaria)和Fc02 (Davidiella)主要分布在叶片和茎中，而Fc32 (Glomus)、F44 (Claroideoglomus)和F58 (Rhizophagus)则主要分布在根中(图2B)。

因此，不同类群的垂直传递内生菌通过繁殖体进入子代植株的不同组织和器官中来发挥其相应的效应。

5 VTE资源的利用

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合理管控植物相关微生物组有望推动第二次绿色革命的到来^[104]。VTE在子代植物相关微生物(包括内生菌、叶际和根际微生物)组装过程中起到引导乃至主导作用，使其成为调控植物微生物的重要切入点。此外，VTE有机会在后代植株中产生持续效应的机制，为通过调控VTE菌群结构来改良植物，进而获得持续稳定的性状提供了可能^[8]。目前，针对VTE的利用仍处于初步阶段。首个利用单一VTE菌株改良植物性状的研究成果来自于葡萄研究^[12]。该研究成功地将葡萄中发掘的一株具有广谱拮抗病原真菌能力的VTE菌株ZX-2导入葡萄扦插枝中，使得该扦插枝萌发的当年及次年植株均获得了相应的抗病功能，并同时表现出一定的促生长和延缓葡萄叶片脱落的性状^[12]。除了直接导入功能型VTE的方式外，通过选育自然获得的有益VTE菌群的繁殖体并进行扩繁，也是VTE利用的一条有效途径。针对VTE的有效开发与利用，期待更多的研究成果和成功案例涌现。

6 总结与展望

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种子及其他植物繁殖体介导了植物内生菌从一代植物到下一代植物的传播。植物在进化过程中选择性地将一些有益VTE长期保持在体内，形成了相对稳定的共生关系^[105]，这是植物适应环境的一种重要机制。目前，有关VTE的研究主要集中于少数植物^{[1,8,12,20,24,106]}，而针对其他植物的VTE资源发掘及其在农业生产中的应用，仍有待更多的研究成果。

种子内生菌可以在子代植物的各个部位，如种子、芽和根中定殖，甚至能够进入根际土壤或植物所处的环境中。反之，种子内生菌也可通过亲代种子、花粉、土壤、大气以及昆虫传播等多种方式获得，这揭示了植物种子获取有益内生菌途径的多样性^[15]。这些与种子紧密相关的微生物群体在种子萌发、幼苗建立以及整体植物适应性方面均发挥着重要作用^[107]，并有可能成为新的VTE成员。VTE在植物中的分布、传递及其与宿主间的相互作用已被广泛研究，揭示了这些微生物如何在植物种群中持续存在并发挥功能^[108]。宿主特性、环境因素、系统发育关系以及地理隔离等因素共同影响着植物内生微生物群落的组装，这展示了植物-微生物-环境之间错综复杂的关系，而深入了解植物VTE的分布特征，对于预测植物群落和生态系统的动态变化也至关重要^[105]。

目前，针对植物VTE的传播规律尚不完全清晰，种子相关微生物群到幼苗的垂直传播途径也尚未完全明确，仍有很大比例的VTE尚未被发掘。此外，影响植物内生菌垂直传递的因素及其作用机制对于多数类群的植物而言仍然难以捉摸。种传VTE存在物种间和个体间的差异性，但对其存在差异的原因研究尚不充分。为了更深入地探究植物种子内生菌的垂直传播机制，可以对多代种子内生菌进行更为系统的跟踪研究，并结合土壤条件及环境因素，综合分析外界环境对植物各组织内生菌群的影响。因此，进一步研究VTE在植物不同部位定殖、扩散以及垂直传递的相关因素，将是今后的研究重点之一。

植物内生菌可以通过种子以及无性繁殖体进行垂直传递，充分探明VTE在宿主中的各种功能，将为这些资源的有效应用开辟更多机会。此外，同一VTE菌株在不同宿主中产生的效应和作用机制可能截然不同。例如，假单胞菌通常被视为某些植物的病原体，但当其作为水稻的垂直传递内生菌时，却具有促进水稻生长和提高水稻抗病性的作用^[48,60]。同样，链格孢属、枝孢属等病原真菌，虽然常造成许多作物产量和品质大量损失，但在芦苇^[63]和香椿^[95]等植物中，却展现出促进植物生长的作用。因此，未来的研究重点应放在深入探究植物体内垂直传递的内生菌如何与宿主植物相互作用，并阐明这些内生菌在不同宿主植物中产生效应差异的内在机制，从而促进植物VTE更加合理和有效的开发利用。

伴随着对植物VTE、植物相关微生物以及它们与宿主间相互作用的深入理解，相关调控技术的不断开发和应用，无疑将迎来植物微生物遗传育种的新时代^[109]。

基金

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国家自然科学基金(32360255)
国家自然科学基金(32471746)
云南省重大科技专项计划(202102AE090042)
云南省科技厅-云南大学 “双一流”建设联合基金(2019FY003024)

参考文献

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文献

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参考文献引证文献

排序方式：

[1]

HARDOIM

, van OVERBEEK

, BERG

, PIRTTILÄ

, COMPANT

, CAMPISANO

, DÖRING

, SESSITSCH

. The hidden world within plants: ecological and evolutionary considerations for defining functioning of microbial endophytes[J]. Microbiology and Molecular Biology Reviews, 2015, 79(3): 293-320.

[2]

SESSITSCH

, REITER

, PFEIFER

, WILHELM

. Cultivation-independent population analysis of bacterial endophytes in three potato varieties based on eubacterial and Actinomycetes-specific PCR of 16S rRNA genes[J]. FEMS Microbiology Ecology, 2002, 39(1): 23-32.

[3]

CHESNEAU

, LAROCHE

, PRÉVEAUX

, MARAIS

, BRIAND

, MAROLLEAU

, SIMONIN

, BARRET

. Single seed microbiota: assembly and transmission from parent plant to seedling[J]. mBio, 2022, 13(6): e0164822.

[4]

BRIGHT

, BULGHERESI

. A complex journey: transmission of microbial symbionts[J]. Nature Reviews Microbiology, 2010, 8(3): 218-230.

[5]

TRUYENS

, WEYENS

, CUYPERS

, VANGRONSVELD

. Bacterial seed endophytes: genera, vertical transmission and interaction with plants[J]. Environmental Microbiology Reports, 2015, 7(1): 40-50.

[6]

HARDOIM

, HARDOIM

CCP

, van OVERBEEK

, van ELSAS

. Dynamics of seed-borne rice endophytes on early plant growth stages[J]. PLoS One, 2012, 7(2): e30438.

[7]

LUO

, TAO

, JUPA

, LIU

, WU

, SONG

, LI

, HUANG

, ZOU

, LIANG

, LI

. Role of vertical transmission of shoot endophytes in root-associated microbiome assembly and heavy metal hyperaccumulation in Sedum alfredii [J]. Environmental Science & Technology, 2019, 53(12): 6954-6963.

[8]

XIANG

, WANG

, CHEN

, LIAO

, LI

, PAN

, ZHU

, YANG

. Dominated “inheritance” of endophytes in grapevines from stock plants via in vitro-cultured plantlets: the dawn of plant endophytic modifications[J]. Horticulturae, 2023, 9(2): 180.

[9]

VANNIER

, MONY

, BITTEBIERE

, MICHON-COUDOUEL

, BIGET

, VANDENKOORNHUYSE

. A microorganisms' journey between plant generations[J]. Microbiome, 2018, 6(1): 79.

[10]

王志伟, 纪燕玲, 陈永敢. 植物内生菌研究及其科学意义[J]. 微生物学通报, 2015, 42(2): 349-363.

WANG

, JI

, CHEN

. Studies and biological significances of plant endophytes[J]. Microbiology China, 2015, 42(2): 349-363 (in Chinese).

[11]

SHADE

, JACQUES

, BARRET

. Ecological patterns of seed microbiome diversity, transmission, and assembly[J]. Current Opinion in Microbiology, 2017, 37: 15-22.

[12]

CHEN

, GUO

, WANG

, WEN

, LI

, LU

, ZHOU

, HUANG

, LI

, PAN

, ZHU

, YANG

. Targeted manipulation of vertically transmitted endophytes to confer beneficial traits in grapevines[J]. Horticulturae, 2024, 10(6): 607.

[13]

ABDELFATTAH

, WISNIEWSKI

, SCHENA

, TACK

AJM

. Experimental evidence of microbial inheritance in plants and transmission routes from seed to phyllosphere and root[J]. Environmental Microbiology, 2021, 23(4): 2199-2214.

[14]

SAMREEN

, NAVEED

, NAZIR

, ASGHAR

, KHAN

, AHMAD ZAHIR

, KANWAL

, JEEVAN

, SHARMA

, MEENA

, SARKAR

, DEVIKA

, PARIHAR

, CHOUDHARY

. Seed associated bacterial and fungal endophytes: diversity, life cycle, transmission, and application potential[J]. Applied Soil Ecology, 2021, 168: 104191.

[15]

FRANK

, SALDIERNA GUZMÁN

, SHAY

. Transmission of bacterial endophytes[J]. Microorganisms, 2017, 5(4): 70.

[16]

FERREIRA

, QUECINE

, LACAVA

, ODA S, AZEVEDO

, ARAÚJO

. Diversity of endophytic bacteria from Eucalyptus species seeds and colonization of seedlings by Pantoea agglomerans [J]. FEMS Microbiology Letters, 2008, 287(1): 8-14.

[17]

GAGNE-BOURGUE

, ALIFERIS

, SEGUIN

, RANI

, SAMSON

, JABAJI

. Isolation and characterization of indigenous endophytic bacteria associated with leaves of switchgrass (Panicum virgatum L.) cultivars[J]. Journal of Applied Microbiology, 2013, 114(3): 836-853.

[18]

COPE-SELBY

, COOKSON

, SQUANCE

, DONNISON

, FLAVELL

, FARRAR

. Endophytic bacteria in Miscanthus seed: implications for germination, vertical inheritance of endophytes, plant evolution and breeding[J]. GCB Bioenergy, 2017, 9(1): 57-77.

[19]

THOMAS

, SHAIK

. Molecular profiling on surface-disinfected tomato seeds reveals high diversity of cultivation-recalcitrant endophytic bacteria with low shares of spore-forming firmicutes[J]. Microbial Ecology, 2020, 79(4): 910-924.

[20]

WALITANG

, KIM

, JEON

, KANG

, SA TM. Conservation and transmission of seed bacterial endophytes across generations following crossbreeding and repeated inbreeding of rice at different geographic locations[J]. MicrobiologyOpen, 2019, 8(3): e00662.

[21]

KAGA

, MANO

, TANAKA

, WATANABE

, KANEKO

, MORISAKI

. Rice seeds as sources of endophytic bacteria[J]. Microbes and Environments, 2009, 24(2): 154-162.

[22]

WANG

, ZHAI

, CAO

, TAN

, ZHANG

. Endophytic bacterial and fungal microbiota in sprouts, roots and stems of rice (Oryza sativa L.)[J]. Microbiological Research, 2016, 188: 1-8.

[23]

ZHANG

, MA

, WANG

, LIAO

, HE

, ZHAO

, GUO

, ZHAO

, WEI

. Dynamics of rice microbiomes reveal core vertically transmitted seed endophytes[J]. Microbiome, 2022, 10(1): 216.

[24]

JOHNSTON-MONJE

, RAIZADA

. Conservation and diversity of seed associated endophytes in Zea across boundaries of evolution, ethnography and ecology[J]. PLoS One, 2011, 6(6): e20396.

[25]

LIU

, ZUO

, XU

, ZOU

, SONG

. Study on diversity of endophytic bacterial communities in seeds of hybrid maize and their parental lines[J]. Archives of Microbiology, 2012, 194(12): 1001-1012.

[26]

RIJAVEC

, LAPANJE

, DERMASTIA

, RUPNIK

. Isolation of bacterial endophytes from germinated maize kernels[J]. Canadian Journal of Microbiology, 2007, 53(6): 802-808.

[27]

梁妍, 宋向东, 谢宗秀, 刘宇, 郝宝成, 王震, 王保海, 梁剑平. 疯草内生真菌研究进展[J]. 动物医学进展, 2020, 41(9): 106-110.

LIANG

, SONG

, XIE

, LIU

, HAO

, WANG

, LIANG

. Progress on endophytic fungi in locoweed[J]. Progress in Veterinary Medicine, 2020, 41(9): 106-110 (in Chinese).

[28]

WHITE

. Widespread distribution of endophytes in the Poaceae [J]. Plant Disease, 1987, 71(4): 340.

[29]

CLAY

, SCHARDL

. Evolutionary origins and ecological consequences of endophyte symbiosis with grasses[J]. The American Naturalist, 2002, 160(): S99-S127.

[30]

MOON

, CRAVEN

, LEUCHTMANN

, CLEMENT

, SCHARDL

. Prevalence of interspecific hybrids amongst asexual fungal endophytes of grasses[J]. Molecular Ecology, 2004, 13(6): 1455-1467.

[31]

TADYCH

, AMBROSE

, BERGEN

, BELANGER

, WHITE JF Jr. Taxonomic placement of Epichloë poae sp. nov. and horizontal dissemination to seedlings via conidia[J]. Fungal Diversity, 2012, 54(1): 117-131.

[32]

赵晓静, 王萍, 李秀璋, 古丽君, 李春杰. 内生真菌在禾草体内的分布特征[J]. 草业科学, 2015, 32(8): 1206-1215.

ZHAO

, WANG

, LI

, GU

, LI

. Distribution characteristics of Epichloë endophyte in gramineous grasses[J]. Pratacultural Science, 2015, 32(8): 1206-1215 (in Chinese).

[33]

IANNONE

, CARGO PD

, GIUSSANI

, SCHARDL

. Geographic distribution patterns of vertically transmitted endophytes in two native grasses in Argentina [J]. Symbiosis, 2013, 59(2): 99-110.

[34]

HODGSON

, de CATES

, HODGSON

, MORLEY

, SUTTON

, GANGE

. Vertical transmission of fungal endophytes is widespread in forbs[J]. Ecology and Evolution, 2014, 4(8): 1199-1208.

[35]

陈泰祥. 野大麦内生真菌提高宿主耐盐性的生理机制研究[D]. 兰州: 兰州大学博士学位论文, 2019.

CHEN

. Study on physiological mechanism of endophytic fungi in wild barley to improve host salt tolerance[D]. Lanzhou: Doctoral Dissertation of Lanzhou University, 2019 (in Chinese).

[36]

GARCÍA PARISI

, CASAS

, GUNDEL

, OMACINI

. Consequences of grazing on the vertical transmission of a fungal Neotyphodium symbiont in an annual grass population[J]. Austral Ecology, 2012, 37(5): 620-628.

[37]

KANG

, JI

, SUN

, ZHAN

, LI

, YU

, WANG

. Taxonomy of Neotyphodium endophytes of Chinese native Roegneria plants[J]. Mycologia, 2009, 101(2): 211-219.

[38]

SELVAKUMAR

, KUNDU

, JOSHI

, NAZIM

, GUPTA

, MISHRA

, GUPTA

. Characterization of a cold-tolerant plant growth-promoting bacterium Pantoea dispersa 1A isolated from a sub-alpine soil in the North Western Indian Himalayas[J]. World Journal of Microbiology and Biotechnology, 2008, 24(7): 955-960.

[39]

MORA

, DÍAZ

, VARGAS-LAGUNAS

, PERALTA

, GUERRERO

, AGUILAR

, ENCARNACIÓN

, GIRARD

, MORA

. Nitrogen-fixing rhizobial strains isolated from common bean seeds: phylogeny, physiology, and genome analysis[J]. Applied and Environmental Microbiology, 2014, 80(18): 5644-5654.

[40]

VERMA

, KINGSLEY

, IRIZARRY

, BERGEN

, KHARWAR

, WHITE JF

. Seed-vectored endophytic bacteria modulate development of rice seedlings[J]. Journal of Applied Microbiology, 2017, 122(6): 1680-1691.

[41]

LOIRET

, ORTEGA

, KLEINER

, ORTEGA-RODÉS

, RODÉS

, DONG

. A putative new endophytic nitrogen-fixing bacterium Pantoea sp. from sugarcane[J]. Journal of Applied Microbiology, 2004, 97(3): 504-511.

[42]

KHALAF

, RAIZADA

. Bacterial seed endophytes of domesticated cucurbits antagonize fungal and oomycete pathogens including powdery mildew[J]. Frontiers in Microbiology, 2018, 9: 42.

[43]

FENG

, SHEN

, SONG

. Rice endophyte Pantoea agglomerans YS19 promotes host plant growth and affects allocations of host photosynthates[J]. Journal of Applied Microbiology, 2006, 100(5): 938-945.

[44]

HAMEED

, YEH

, HSIEH

, CHUNG

, LO CT, YOUNG

. Diversity and functional characterization of bacterial endophytes dwelling in various rice (Oryza sativa L.) tissues, and their seed-borne dissemination into rhizosphere under gnotobiotic P-stress[J]. Plant and Soil, 2015, 394(1/2): 177-197.

[45]

TSAVKELOVA

, CHERDYNTSEVA

, BOTINA

, NETRUSOV

. Bacteria associated with orchid roots and microbial production of auxin[J]. Microbiological Research, 2007, 162(1): 69-76.

[46]

JASIM

, JOHN JIMTHA

, JYOTHIS

, RADHAKRISHNAN

. Plant growth promoting potential of endophytic bacteria isolated from Piper nigrum [J]. Plant Growth Regulation, 2013, 71(1): 9802.

[47]

TSEGAYE

, ASSEFA

, TEFERA

, ALEMU

, GIZAW

. Characterization and identification of tef (Eragrostis tef) seed endophytic bacterial species and evaluate their effect on plant growth promotion[J]. Journal of Plant Pathology & Microbiology, 2018, 9(4): 438.

[48]

WANG

, ZHAI

, CAO

, TAN

, ZHANG

. Illumina-based analysis of core actinobacteriome in roots, stems, and grains of rice[J]. Microbiological Research, 2016, 190: 12-18.

[49]

曹艳花, 徐凤花, 陈小忠, 张晓霞, 韦善君, 马晓彤. 水稻种子相关细菌的系统发育分析与促生能力评价[J]. 中国土壤与肥料, 2011(5): 83-87.

CAO

, XU

, CHEN

, ZHANG

, WEI

, MA

. Analysis on growth-promoting ability and phylogeny with rice seed associated bacteria[J]. Soil and Fertilizer Sciences in China, 2011(5): 83-87 (in Chinese).

[50]

PUENTE

, LI

, BASHAN

. Endophytic bacteria in cacti seeds can improve the development of cactus seedlings[J]. Environmental and Experimental Botany, 2009, 66(3): 402-408.

[51]

罗光明, 董艳凯, 朱玉野, 朱继孝, 曾金祥, 王晓云, 胡燕珍, 罗扬婧, 龚雨虹. 栀子种子内生菌检测及药剂消毒处理效果比较研究[J]. 中国药学杂志, 2015, 50(19): 1665-1669.

LUO

, DONG

, ZHU

, ZENG

, WANG

, HU

, LUO

, GONG

. Detection of endogenous bacteria of Gardenia seed and comparative study of chemical treatments[J]. China Industrial Economics, 2015, 50(19): 1665-1669 (in Chinese).

[52]

冯玮娜, 彭培好. 四川牡丹根际微生物及种子内生菌组成[J]. 东北林业大学学报, 2020, 48(1): 88-94.

FENG

, PENG

. Microbial composition associated with the rhizosphere and seed endosphere of Paeonia szechuanica [J]. Journal of Northeast Forestry University, 2020, 48(1): 88-94 (in Chinese).

[53]

, ZHOU

, LIU

. Diversity of endophytic bacteria in Cardamine hupingshanensis and potential of culturable selenium-resistant endophytes to enhance seed germination under selenate stress[J]. Current Microbiology, 2021, 78(5): 2091-2103.

[54]

CHOI

, JEONG

, KIM

. Variation of the seed endophytic bacteria among plant populations and their plant growth-promoting activities in a wild mustard plant species, Capsella bursa-pastoris [J]. Ecology and Evolution, 2022, 12(3): e8683.

[55]

KRISHNAMOORTHY

, AGARWAL

, KOTAMREDDY

JNR

, BHATTACHARYA

, MITRA

, MAITI

. Impact of seed-transmitted endophytic bacteria on intra- and inter-cultivar plant growth promotion modulated by certain sets of metabolites in rice crop[J]. Microbiological Research, 2020, 241: 126582.

[56]

SCOTT

, RANI

, SAMSATLY

, CHARRON

, JABAJI

. Endophytes of industrial hemp (Cannabis sativa L.) cultivars: identification of culturable bacteria and fungi in leaves, petioles, and seeds[J]. Canadian Journal of Microbiology, 2018, 64(10): 664-680.

[57]

PUENTE

, LI

, BASHAN

. Microbial populations and activities in the rhizoplane of rock-weathering desert plants. II. growth promotion of cactus seedlings[J]. Plant Biology, 2004, 6(5): 643-650.

[58]

VERMA

, WHITE

. Indigenous endophytic seed bacteria promote seedling development and defend against fungal disease in browntop millet (Urochloa ramosa L.)[J]. Journal of Applied Microbiology, 2018, 124(3): 764-778.

[59]

KIM

, ROY

, AHN SH, SHANMUGAM

, YANG

, JUNG

, JEON

. Culturable endophytes associated with soybean seeds and their potential for suppressing seed-borne pathogens[J]. The Plant Pathology Journal, 2022, 38(4): 313-322.

[60]

WHITE

, KINGSLEY

, KOWALSKI

, IRIZARRY

, MICCI

, SOARES

, BERGEN

. Disease protection and allelopathic interactions of seed-transmitted endophytic pseudomonads of invasive reed grass (Phragmites australis)[J]. Plant and Soil, 2018, 422(1/2): 195-208.

[61]

DÍAZ HERRERA

, GROSSI

, ZAWOZNIK

, GROPPA

. Wheat seeds harbour bacterial endophytes with potential as plant growth promoters and biocontrol agents of Fusarium graminearum [J]. Microbiological Research, 2016, 186: 37-43.

[62]

FÜRNKRANZ

, LUKESCH

, MÜLLER

, HUSS

, GRUBE

, BERG

. Microbial diversity inside pumpkins: microhabitat-specific communities display a high antagonistic potential against phytopathogens[J]. Microbial Ecology, 2012, 63(2): 418-428.

[63]

SHEARIN

ZRC

, FILIPEK

, DESAI

, BICKFORD

, KOWALSKI

, CLAY

. Fungal endophytes from seeds of invasive, non-native Phragmites australis and their potential role in germination and seedling growth[J]. Plant and Soil, 2018, 422(1): 183-194.

[64]

LIDSTROM

, CHISTOSERDOVA

. Plants in the pink: cytokinin production by Methylobacterium [J]. Journal of Bacteriology, 2002, 184(7): 1818.

[65]

GOGGIN

, EMERY

, KUREPIN

, POWLES

. A potential role for endogenous microflora in dormancy release, cytokinin metabolism and the response to fluridone in Lolium rigidum seeds[J]. Annals of Botany, 2015, 115(2): 293-301.

[66]

ALIBRANDI

, CARDINALE

, RAHMAN

, STRATI

, CINÁ

, de VIANA

, GIAMMINOLA

, GALLO

, SCHNELL

, de FILIPPO

, CIACCIO

, PUGLIA

. The seed endosphere of Anadenanthera colubrina is inhabited by a complex microbiota, including Methylobacterium spp. and Staphylococcus spp. with potential plant-growth promoting activities[J]. Plant and Soil, 2018, 422(1/2): 81-99.

[67]

SÁNCHEZ-LÓPEZ

, PINTELON

, STEVENS

, IMPERATO

, TIMMERMANS

, GONZÁLEZ-CHÁVEZ

, CARRILLO-GONZÁLEZ

, van HAMME

, VANGRONSVELD

, THIJS

. Seed endophyte microbiome of Crotalaria pumila unpeeled: identification of plant-beneficial methylobacteria[J]. International Journal of Molecular Sciences, 2018, 19(1): 291.

[68]

艾加敏. 白刺花根瘤中非根瘤菌与根瘤菌组装、演替及其互作关系研究[D]. 延安: 延安大学硕士学位论文, 2023.

. Assembly, succession and interaction between non-rhizobium and rhizobium in nodules of Sophora davidii (Franch.) Skeels[D]. Yan'an: Master's Thesis of Yan'an University, 2023 (in Chinese).

[69]

GOND

, BERGEN

, TORRES

, WHITE JF

. Endophytic Bacillus spp. produce antifungal lipopeptides and induce host defence gene expression in maize[J]. Microbiological Research, 2015, 172: 79-87.

[70]

JOOSTE

, ROETS

, MIDGLEY

, OBERLANDER

, DREYER

. Nitrogen-fixing bacteria and Oxalis-evidence for a vertically inherited bacterial symbiosis[J]. BMC Plant Biology, 2019, 19(1): 441.

[71]

ZHU

. Isolation and identification of Ammodendron bifolium endophytic bacteria and the action mechanism of selected isolates-induced seed germination and their effects on host osmotic-stress tolerance[J]. Archives of Microbiology, 2019, 201(4): 431-442.

[72]

, SHENG

, CHEN

, MEN YJ, GAN

, GUO

, SHEN

. Bacterial community compositions of tomato (Lycopersicum esculentum Mill.) seeds and plant growth promoting activity of ACC deaminase producing Bacillus subtilis (HYT-12-1) on tomato seedlings[J]. World Journal of Microbiology & Biotechnology, 2014, 30(3): 835-845.

[73]

CHOI

, KIM

, JEONG

, KIM

. Effects of seed endophytic bacteria on life history and reproductive traits in a cosmopolitan weed, Capsella Bursa-pastoris [J]. Plants, 2022, 11(19): 2642.

[74]

SHAHZAD

, WAQAS

, KHAN

, ASAF

, KHAN

, KANG

, YUN

, LEE

. Seed-borne endophytic Bacillus amyloliquefaciens RWL-1 produces gibberellins and regulates endogenous phytohormones of Oryza sativa [J]. Plant Physiology and Biochemistry, 2016, 106: 236-243.

[75]

颜瑾, 舒翠华, 田昌, 王运生, 肖启明, 陈武, 巢进. 烤烟K326种子可培养内生细菌的分离与鉴定[J]. 湖南农业科学, 2014(18): 21-24, 27.

YAN

, SHU

, TIAN

, WANG

, XIAO

, CHEN

, CHAO

. Isolation and identification of endophytic bacteria in seeds of tobacco variety K326[J]. Hunan Agricultural Sciences, 2014(18): 21-24, 27 (in Chinese).

[76]

DOWARAH

, AGARWAL

, KRISHNATREYA

, SHARMA

, KALITA

, AGARWALA

. Evaluation of seed associated endophytic bacteria from tolerant chilli cv. Firingi Jolokia for their biocontrol potential against bacterial wilt disease[J]. Microbiological Research, 2021, 248: 126751.

[77]

MARZOUK

, CHAOUACHI

, SHARMA

, JALLOULI

, MHAMDI

, KAUSHIK

, DJÉBALI

. Biocontrol of Rhizoctonia solani using volatile organic compounds of solanaceae seed-borne endophytic bacteria[J]. Postharvest Biology and Technology, 2021, 181: 111655.

[78]

MOUSA

, SHEARER

, LIMAY-RIOS

, ZHOU

, RAIZADA

. Bacterial endophytes from wild maize suppress Fusarium graminearum in modern maize and inhibit mycotoxin accumulation[J]. Frontiers in Plant Science, 2015, 6: 805.

[79]

YANG

, ZHANG

, WU

, XU

, AHMAD

, ZHANG

, ZHAO

, LIU

. An endophytic strain of the genus Bacillus isolated from the seeds of maize (Zea mays L.) has antagonistic activity against maize pathogenic strains[J]. Microbial Pathogenesis, 2020, 142: 104074.

[80]

蔡学清, 何红, 胡方平. 内生菌BS-2对辣椒苗的促生作用及对内源激素的影响[J]. 亚热带农业研究, 2005, 1(4): 49-52.

CAI

, HE

, HU

. The effects of endophytic BS-2 (Bacillus subtilis) on growth and internal phytohormone of Capsicum [J]. Subtropical Agriculture Research, 2005, 1(4): 49-52 (in Chinese).

[81]

李雨轩. 紫茎泽兰叶组织和冠层空气真菌时空动态及其种子内生真菌研究[D]. 昆明: 云南大学硕士学位论文, 2021.

. Temporal and spatial dynamics of the fungi in leaf tissues and canopy air of Aseratina adenophora and its seed endophytic fungi[D]. Kunming: Master's Thesis of Yunnan University, 2021 (in Chinese).

[82]

RAHMAN

, FLORY

, KOYRO

, ABIDEEN

, SCHIKORA

, SUAREZ

, SCHNELL

, CARDINALE

. Consistent associations with beneficial bacteria in the seed endosphere of barley (Hordeum vulgare L.)[J]. Systematic and Applied Microbiology, 2018, 41(4): 386-398.

[83]

SULLIVAN

, ROBERTS

, BULTMAN

. Genetic covariation between the vertically transmitted endophyte Epichloë canadensis and its host Canada wildrye[J]. Microbial Ecology, 2023, 86(3): 1686-1695.

[84]

ROZPĄDEK

, WĘŻOWICZ

, NOSEK

, WAŻNY

, TOKARZ

, LEMBICZ

, MISZALSKI

, TURNAU

. The fungal endophyte Epichloë typhina improves photosynthesis efficiency of its host orchard grass (Dactylis glomerata)[J]. Planta, 2015, 242(4): 1025-1035.

[85]

ELBERSEN

, WEST

. Growth and water relations of field-grown tall fescue as influenced by drought and endophyte[J]. Grass and Forage Science, 1996, 51(4): 333-342.

[86]

KANE

. Effects of endophyte infection on drought stress tolerance of Lolium perenne accessions from the Mediterranean region[J]. Environmental and Experimental Botany, 2011, 71(3): 337-344.

[87]

AIKEN

, STRICKLAND

. Forages and Pastures Symposium: managing the tall fescue-fungal endophyte symbiosis for optimum forage-animal production[J]. Journal of Animal Science, 2013, 91(5): 2369-2378.

[88]

PENNELL

, ROLSTON

, de BONTH

, SIMPSON

, DE

. Development of a bird-deterrent fungal endophyte in turf tall fescue[J]. New Zealand Journal of Agricultural Research, 2010, 53(2): 145-150.

[89]

LEFORT

, McKINNON

, NELSON

, GLARE

. Natural occurrence of the entomopathogenic fungi Beauveria bassiana as a vertically transmitted endophyte of Pinus radiata and its effect on above and belowground insect pests[J]. New Zealand Plant Protection, 2016, 69: 68-77.

[90]

SHI

. Alfalfa endogenous rhizobia[J]. Agricultural Research & Technology, 2018, 13(4): 1614-1623.

[91]

康文娟, 师尚礼, 王泽一, 陈建纲, 苗阳阳. 3株紫花苜蓿内生根瘤菌功能差异性分析[J]. 草业科学, 2018, 35(7): 1614-1623.

KANG

, SHI

, WANG

, CHEN

, MIAO

. Analysis of functional differences among three Medicago sativa endophytic rhizobial strains[J]. Pratacultural Science, 2018, 35(7): 1614-1623 (in Chinese).

[92]

祁娟, 师尚礼. 不同品种紫花苜蓿种子内生根瘤菌溶磷和分泌生长素能力[J]. 草原与草坪, 2006, 26(5): 18-20, 25.

, SHI

. Preliminary study on the ability of phosphorus-solubilizing and IAA-secreting of endogenous rhizobia in seeds of different alfalfa varieties[J]. Grassland and Turf, 2006, 26(5): 18-20, 25 (in Chinese).

[93]

TRUYENS

, BECKERS

, THIJS

, WEYENS

, CUYPERS

, VANGRONSVELD

. Cadmium-induced and trans-generational changes in the cultivable and total seed endophytic community of Arabidopsis thaliana [J]. Plant Biology, 2016, 18(3): 376-381.

[94]

WIJESOORIYA

WADK

, DESHAPPRIYA

. An inoculum of endophytic fungi for improved growth of a traditional ricevariety in Sri Lanka [J]. Tropical Plant Research, 2016, 3(3): 470-480.

[95]

RAHMAWATI

, ISFANDITO

, ASTUTI

, ADITIAWATI

. Endophytic fungi from Surian (Toona sinensis Roem) and antioxidant potency from its culture[J]. Asian Journal of Plant Sciences, 2015, 15(1/2): 8-15.

[96]

RASHID

, CHARLES

, GLICK

. Isolation and characterization of new plant growth-promoting bacterial endophytes[J]. Applied Soil Ecology, 2012, 61: 217-224.

[97]

RUIZA

, AGARAS

, de WERRAB

, WALL

, VALVERDE

. Characterization and screening of plant probiotic traits of bacteria isolated from rice seeds cultivated in Argentina [J]. The Journal of Microbiology, 2011, 49(6): 902-912.

[98]

李秀璋. 醉马草内生真菌与宿主种带真菌、根际微生物的互作及其进化研究[D]. 兰州: 兰州大学博士学位论文, 2017.

. Study on the interaction and evolution of endophytic fungi, host species-bearing fungi and rhizosphere microorganisms in Buddleja [D]. Lanzhou: Doctoral Dissertation of Lanzhou University, 2017 (in Chinese).

[99]

朱文勇, 李洁, 赵国振, 黄海玉, 秦盛, 赵立兴, 徐丽华. 喜树内生放线菌多样性及抗菌活性评价[J]. 微生物学通报, 2010, 37(2): 211-216.

ZHU

, LI

, ZHAO

, HUANG

, QIN

, ZHAO

, XU

. Diversity and antimicrobial activities of endophytic actinomycetes isolated from Camptotheca acuminata Decne.[J]. Microbiology China, 2010, 37(2): 211-216 (in Chinese).

[100]

姚晓玲, 康前进, 熊顺子, 李芳, 王益, 林双君, 白林泉, 马伟, 邓子新. 喜树种子内生放线菌的分离鉴定及抗菌活性物质初分离[J]. 微生物学通报, 2014, 41(6): 1109-1120.

YAO

, KANG

, XIONG

, LI

, WANG

, LIN

, BAI

, MA

, DENG

. Isolation and identification of endophytic actinomycetes from the seeds of Camptotheca acuminata Decne. and isolation of antimicrobial substances from those endophytic actinomycetes[J]. Microbiology China, 2014, 41(6): 1109-1120 (in Chinese).

[101]

赵霞. 水稻种子内生细菌多样性分析及核心微生物组的界定[D]. 北京: 中国农业科学院硕士学位论文, 2019.

ZHAO

. Diversity analysis of endophytic bacteria in rice seeds and definition of core microbial groups[D]. Beijing: Master's Thesis of Chinese Academy of Agricultural Sciences, 2019 (in Chinese).

[102]

COMPANT

, REITER

, SESSITSCH

, NOWAK

, CLÉMENT

, BARKA E

AIT

. Endophytic colonization of Vitis vinifera L. by plant growth-promoting bacterium Burkholderia sp. strain PsJN[J]. Applied and Environmental Microbiology, 2005, 71(4): 1685-1693.

[103]

王美宁. 中华羊茅内生真菌培养特性及其金属离子抗性的研究[D]. 兰州: 兰州大学硕士学位论文, 2019.

WANG

. Study on culture characteristics and metal ion resistance of endophytic fungi from Festuca sinensis [D]. Lanzhou: Master's Thesis of Lanzhou University, 2019 (in Chinese).

[104]

, ZHENG

, YADAV

, SAHA

, SALAMA

, LI

, WANG

, JEON

. Rational management of the plant microbiome for the Second Green Revolution[J]. Plant Communications, 2024, 5(4): 100812.

[105]

RUDGERS

, AFKHAMI

, RÚA

, DAVITT

, HAMMER

, HUGUET

. A fungus among us: broad patterns of endophyte distribution in the grasses[J]. Ecology, 2009, 90(6): 1531-1539.

[106]

COMPANT

, CLÉMENT

, SESSITSCH

. Plant growth-promoting bacteria in the rhizo- and endosphere of plants: their role, colonization, mechanisms involved and prospects for utilization[J]. Soil Biology and Biochemistry, 2010, 42(5): 669-678.

[107]

SEMENZATO

, FADDETTA

, FALSINI

, del DUCA

, ESPOSITO

, PADULA

, GRECO

, MUCCI

, ZACCARONI

, PUGLIA

, PAPINI

, FANI

. Endophytic bacteria associated with Origanum heracleoticum L. (Lamiaceae) seeds[J]. Microorganisms, 2022, 10(10): 2086.

[108]

GIBERT

, MAGDA

, HAZARD

. Interplay between endophyte prevalence, effects and transmission: insights from a natural grass population[J]. PLoS One, 2015, 10(10): e0139919.

[109]

CERNAVA

. Coming of age for microbiome gene breeding in plants[J]. Nature Communications, 2024, 15(1): 6623.

2025年第65卷第2期

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doi: 10.13343/j.cnki.wsxb.20240527

接收时间：2024-08-26
首发时间：2026-02-05
出版时间：2025-02-04

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收稿日期：2024-08-26
录用日期：2024-10-23

基金

National Natural Science Foundation of China(32360255)

国家自然科学基金(32360255)

National Natural Science Foundation of China(32471746)

国家自然科学基金(32471746)

Yunnan Provincial Key Science and Technology Special Project(202102AE090042)

云南省重大科技专项计划(202102AE090042)

Joint Fund for “Double First Class”Construction between Yunnan Provincial Department of Science and Technology and Yunnan University(2019FY003024)

云南省科技厅-云南大学 “双一流”建设联合基金(2019FY003024)

作者信息

1 云南大学生态与环境学院，云南高原山地生态与退化环境修复重点实验室，云南昆明

2 云南农业大学植物保护学院，云南昆明

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2种不同金属材料的力学参数

科 Family	属数 Number of genus	种数 Number of species	占总种数比例 Percentage of total species (%)	属 Genus	种数 Number of species	占总种数比例 Percentage of total species (%)
鹅膏菌科Amanitaceae	2	11	5.26	鹅膏菌属 Amanita	10	4.78
小菇科 Mycenaceae	2	12	5.74	丝盖伞属 Inocybe	5	2.39
多孔菌科 Polyporaceae	8	14	6.70	蜡蘑属 Laccaria	5	2.39
红菇科 Russulaceae	3	23	11.00	小皮伞属 Marasmius	6	2.87
				小菇属 Mycena	11	5.26
				光柄菇属 Pluteus	5	2.39
				红菇属 Russula	17	8.13
				栓菌属 Trametes	5	2.39

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内生菌 Endophyte	VTE属 VTE genus	宿主植物 Plant	载体 Carrier	功能 Functions
内生细菌 Endophytic bacteria	泛菌属Pantoea	水稻Oryza sativa^[23] 玉米Zea mays^[24] 桉树Eucalyptus sp.^[16]	种子Seed 种子Seed 种子Seed	调节植物激素Regulate plant hormones^[38-40] 固氮作用Nitrogen fixation^[41] 产铁载体Produce siderophores^[38-39,41] 增强植物抗病性Enhance plant disease resistance^[42] 促进植物生长Promote plant growth^{[16,23,38-40,43]}
黄单胞菌Xanthomonas	水稻O. sativa^[23] 东南景天Sedum alfredii^[7]	种子Seed 扦插枝Cutting	固氮作用Nitrogen fixation^[23,44] 产铁载体Produce siderophores^[44] 促进植物生长Promote plant growth^[23]
肠杆菌属Enterobacter	水稻O. sativa^[40]	种子Seed	调节植物激素Regulate plant hormones^[40,45-46] 溶解磷能力Phosphorus solubilizing ability^[47] 促进植物生长Promote plant growth^[24,40]
假单胞菌属Pseudomonas	水稻O. sativa^[48]	种子Seed	调节植物激素Regulate plant hormones^[49] 固氮作用Nitrogen fixation^[50-52] 产铁载体Produce siderophores^{[47,49,53-54]} 溶解磷能力Phosphorus solubilizing ability^[55-58] 增强植物抗病性Enhance plant disease resistance^[59-60]
鞘氨醇单胞菌属Sphingomonas	水稻O. sativa^[48] 葡萄Vitis vinifera^[8]	种子Seed 茎尖Shoots	-
新鞘氨醇杆菌属Novosphingobium	葡萄V. vinifera^[8]	茎尖Shoots	-
微杆菌属Microbacterium	水稻O. sativa^[20] 柳枝稷Panicum virgatum^[17]	种子Seed 种子Seed	-
类芽孢杆菌属Paenibacillus	桉树Eucalyptus sp.^[16] 玉米Z. mays^[24]	种子Seed 种子Seed	促进植物生长Promote plant growth^[16,61] 增强植物抗病性Enhance plant disease resistance^[26,62-63]
腐殖质类芽孢杆菌Paenibacillus humicus	桉树Eucalyptus sp.^[16]	种子Seed	-
蒙氏肠球菌Enterococcus mundtii	桉树Eucalyptus sp.^[16]	种子Seed	-
甲基杆菌属Methylobacterium	桉树Eucalyptus sp.^[16]	种子Seed	促进植物生长Promote plant growth^[64-67] 提高重金属耐受Enhance heavy metal tolerance^[67]
梭菌属Clostridium	玉米Z. mays^[24]	种子Seed	-
中生根瘤菌属Mesorhizobium	白刺花Sophora davidii^[68]	种子Seed	增强植物抗逆性Enhance plant stress resistance^[68]
贪铜菌属Cupriavidus	葡萄V. vinifera^[8]	茎尖Shoots	-
劳尔氏菌属Ralstonia	葡萄V. vinifera^[8]	茎尖Shoots	-
芽孢杆菌属Bacillus	水稻O. sativa^[6] 桉树Eucalyptus sp.^[16] 柳枝稷P. virgatum^[17] 仙人掌Opuntia stricta^[50] 番茄Solanum lycopersicum^[69] 酢浆草Oxalis corniculate^[70] 葡萄V. vinifera^[1,8,12]	种子Seed 种子Seed 种子Seed 种子Seed 种子Seed 种子Seed 种子Seed, 扦插枝Cutting	调节植物激素Regulate plant hormones^[71-74] 固氮作用Nitrogen fixation^[50,72] 产铁载体Produce siderophores^[56,75] 溶解磷能力^[53,56-57] 增强植物抗病性Enhance plant disease resistance^{[39,69,76-79]} 促进植物生长Promote plant growth^[39,72,80]
内生真菌 Endophytic fungi	链格孢属Alternaria	欧洲千里光Senecio vulgaris^[34] 长叶车前Plantago lanceolata^[34] 酸模Rumex acetosa^[34] 紫茎泽兰Ageratina adenophora^[81] 葡萄V. vinifera^[8]	种子Seed 种子Seed 种子Seed 种子Seed 茎尖Shoots	促进植物生长Promote plant growth^[63]
Epichloё (Neotyphodium)	禾本科Poaceae^[82] 加拿大黑麦Elymus canadensis^[83] 醉马草Achnatherum inebrian^[35] 野大麦Hordeum brevisubulatum^[35] 高羊茅Festuca arundinacea^[35] 黑麦草Lolium multiflorum^[36] Poa lanuginosa poir^[33] Poe bonariensis^[33]	种子Seed 种子Seed 种子Seed 种子Seed 种子Seed 种子Seed 种子Seed 种子Seed	促进植物生长Promote plant growth^[41,84-85] 改善植物光合作用Improve plant photosynthesis^[41,84] 增强植物抗逆性Enhance plant stress resistance^[86-87] 生产麦角碱和喹啉生物碱Produce ergot alkaloids and quinoline alkaloids^[88]
白僵菌属Beauveria	辐射松Pinus radiata^[89]	种子Seed	-
白粉菌属Erysiphe	葡萄V. vinifera^[8]	茎尖Shoots	-
根瘤菌Rhizosphaera	东南景天S. alfredii^[7] 葡萄V. vinifera^[8]	扦插枝Cutting 茎尖Shoots	固氮作用Nitrogen fixation^[39,90-91] 溶解磷能力Phosphorus solubilizing ability^[47,92] 增强植物抗逆性Enhance plant stress resistance^[92-93]
Davidiella	葡萄V. vinifera^[8]	茎尖Shoots	-
隐球菌属Cryptococcu	紫茎泽兰A. adenophora^[81]	种子Seed	-
枝孢属Cladosporiu	矢车菊Centaurea Cyanus^[34] 罂粟Papaver rhoeas^[34] 黑矢车菊Centaurea nigra^[34] 紫茎泽兰A. adenophora^[81] 葡萄V. vinifera^[8]	种子Seed 种子Seed 种子Seed 种子Seed 茎尖Shoots	促进植物生长Promote plant growth^[94] 抗氧化能力Antioxidant capacity^[95]
内生放线菌 Endophytic actinomycetes	链霉菌属Streptomyces	东南景天S. alfredii^[7]	扦插枝Cutting	促进植物生长Promote plant growth^[7] 增强植物抗逆性Enhance plant stress resistance^[7]
类诺卡氏属Nocardiopsis	东南景天S. alfredii^[7]	扦插枝Cutting	-
假诺卡氏菌属Pseudonocardia	东南景天S. alfredii^[7]	扦插枝Cutting	-

内生菌

Endophyte

VTE属

VTE genus

宿主植物

Plant

载体

Carrier

功能

Functions

内生细菌

Endophytic bacteria

泛菌属Pantoea

水稻Oryza sativa^[23]

玉米Zea mays^[24]

桉树Eucalyptus sp.^[16]

种子Seed

调节植物激素Regulate plant hormones^[38-40]

固氮作用Nitrogen fixation^[41]

产铁载体Produce siderophores^[38-39,41]

增强植物抗病性Enhance plant disease resistance^[42]

促进植物生长Promote plant growth^{[16,23,38-40,43]}

黄单胞菌Xanthomonas

水稻O. sativa^[23]

东南景天Sedum alfredii^[7]

种子Seed

扦插枝Cutting

固氮作用Nitrogen fixation^[23,44]

产铁载体Produce siderophores^[44]

促进植物生长Promote plant growth^[23]

肠杆菌属Enterobacter

水稻O. sativa^[40]

种子Seed

调节植物激素Regulate plant hormones^[40,45-46]

溶解磷能力Phosphorus solubilizing ability^[47]

促进植物生长Promote plant growth^[24,40]

假单胞菌属Pseudomonas

水稻O. sativa^[48]

种子Seed

调节植物激素Regulate plant hormones^[49]

固氮作用Nitrogen fixation^[50-52]

产铁载体Produce siderophores^{[47,49,53-54]}

溶解磷能力Phosphorus solubilizing ability^[55-58]

增强植物抗病性Enhance plant disease resistance^[59-60]

鞘氨醇单胞菌属Sphingomonas

水稻O. sativa^[48]

葡萄Vitis vinifera^[8]

种子Seed

茎尖Shoots

新鞘氨醇杆菌属Novosphingobium

葡萄V. vinifera^[8]

茎尖Shoots

微杆菌属Microbacterium

水稻O. sativa^[20]

柳枝稷Panicum virgatum^[17]

种子Seed

类芽孢杆菌属Paenibacillus

桉树Eucalyptus sp.^[16]

玉米Z. mays^[24]

种子Seed

促进植物生长Promote plant growth^[16,61]

增强植物抗病性Enhance plant disease resistance^[26,62-63]

腐殖质类芽孢杆菌Paenibacillus humicus

桉树Eucalyptus sp.^[16]

种子Seed

蒙氏肠球菌Enterococcus mundtii

桉树Eucalyptus sp.^[16]

种子Seed

甲基杆菌属Methylobacterium

桉树Eucalyptus sp.^[16]

种子Seed

促进植物生长Promote plant growth^[64-67]

提高重金属耐受Enhance heavy metal tolerance^[67]

梭菌属Clostridium

玉米Z. mays^[24]

种子Seed

中生根瘤菌属Mesorhizobium

白刺花Sophora davidii^[68]

种子Seed

增强植物抗逆性Enhance plant stress resistance^[68]

贪铜菌属Cupriavidus

葡萄V. vinifera^[8]

茎尖Shoots

劳尔氏菌属Ralstonia

葡萄V. vinifera^[8]

茎尖Shoots

芽孢杆菌属Bacillus

水稻O. sativa^[6]

桉树Eucalyptus sp.^[16]

柳枝稷P. virgatum^[17]

仙人掌Opuntia stricta^[50]

番茄Solanum lycopersicum^[69]

酢浆草Oxalis corniculate^[70]

葡萄V. vinifera^[1,8,12]

种子Seed

种子Seed,

扦插枝Cutting

调节植物激素Regulate plant hormones^[71-74]

固氮作用Nitrogen fixation^[50,72]

产铁载体Produce siderophores^[56,75]

溶解磷能力^[53,56-57]

增强植物抗病性Enhance plant disease resistance^{[39,69,76-79]}

促进植物生长Promote plant growth^[39,72,80]

内生真菌

Endophytic fungi

链格孢属Alternaria

欧洲千里光Senecio vulgaris^[34]

长叶车前Plantago lanceolata^[34]

酸模Rumex acetosa^[34]

紫茎泽兰Ageratina adenophora^[81]

葡萄V. vinifera^[8]

种子Seed

茎尖Shoots

促进植物生长Promote plant growth^[63]

Epichloё

(Neotyphodium)

禾本科Poaceae^[82]

加拿大黑麦Elymus canadensis^[83]

醉马草Achnatherum inebrian^[35]

野大麦Hordeum brevisubulatum^[35]

高羊茅Festuca arundinacea^[35]

黑麦草Lolium multiflorum^[36]

Poa lanuginosa poir^[33]

Poe bonariensis^[33]

种子Seed

促进植物生长Promote plant growth^[41,84-85]

改善植物光合作用Improve plant photosynthesis^[41,84]

增强植物抗逆性Enhance plant stress resistance^[86-87]

生产麦角碱和喹啉生物碱Produce ergot alkaloids and quinoline alkaloids^[88]

白僵菌属Beauveria

辐射松Pinus radiata^[89]

种子Seed

白粉菌属Erysiphe

葡萄V. vinifera^[8]

茎尖Shoots

根瘤菌Rhizosphaera

东南景天S. alfredii^[7]

葡萄V. vinifera^[8]

扦插枝Cutting

茎尖Shoots

固氮作用Nitrogen fixation^[39,90-91]

溶解磷能力Phosphorus solubilizing ability^[47,92]

增强植物抗逆性Enhance plant stress resistance^[92-93]

Davidiella

葡萄V. vinifera^[8]

茎尖Shoots

隐球菌属Cryptococcu

紫茎泽兰A. adenophora^[81]

种子Seed

枝孢属Cladosporiu

矢车菊Centaurea Cyanus^[34]

罂粟Papaver rhoeas^[34]

黑矢车菊Centaurea nigra^[34]

紫茎泽兰A. adenophora^[81]

葡萄V. vinifera^[8]

种子Seed

茎尖Shoots

促进植物生长Promote plant growth^[94]

抗氧化能力Antioxidant capacity^[95]

内生放线菌

Endophytic actinomycetes

链霉菌属Streptomyces

东南景天S. alfredii^[7]

扦插枝Cutting

促进植物生长Promote plant growth^[7]

增强植物抗逆性Enhance plant stress resistance^[7]

类诺卡氏属Nocardiopsis

东南景天S. alfredii^[7]

扦插枝Cutting

假诺卡氏菌属Pseudonocardia

东南景天S. alfredii^[7]

扦插枝Cutting