Effects of Epichloë endophytic fungi on the germination of grass seeds

doi:10.11686/cyxb2020601

Abstract

Abstract:

Seeds are the most basic output generated in agricultural production， and seed germination plays a decisive role in survival and reproduction of species. However， the process of germination is easily affected by various environment factors. It is significant to improve germination quality in stress condition for grass planting. The establishment of the symbiotic relationship in plants with Epichlo? endophyte confers on the symbiosis higher germination rate， stronger resistance to adverse stress， and leads to the occupation of a more favourable ecological niche in the phytocoenosis. Hence， the Epichlo? endophyte has a wide impact in ecosystems， with ramifications affecting animals， plants and microorganisms within an ecosystem. This study reviewed the influence of Epichlo? endophyte infection on host grass germination under different habitats and treatments， with a focus on roles of endophyte in seed germination under natural growth conditions and various stress conditions， including allelopathy. It was found that endophyte promotes seed germination under both natural and stress conditions. However， germination responses were inconsistent in the presence of allelopathic effects. Additionally， the negative effect on germination caused by extended storage time is alleviated by endophyte infection. Building on previous research， we should prioritize research into the physiology and biochemistry of promoting germination and enhancing resistance to biotic and abiotic stresses in grass infected with Epichlo? endophyte， and on defining the mechanism whereby endophyte fungi influence germination， in order to identify excellent endophytic strains and explore novel ways of creating grass germplasm resources using Epichlo? endophyte.

Key words: grass endophyte, seed germination, biotic and abiotic stress, allelopathy

Chun-jie LI, Ming-xiao LANG, Zhen-jiang CHEN, Tai-xiang CHEN, Jing LIU, Yuan-yuan JIN, Xue-kai WEI. Effects of Epichloë endophytic fungi on the germination of grass seeds[J]. Acta Prataculturae Sinica, 2022, 31(3): 192-206.

Figures/Tables 3

References 97

1	Schardl C L， Leuchtmann A， Spiering M J. Symbioses of grasses with seedborne fungal endophytes. Annual Review of Plant Biology， 2004， 55（1）： 315-340.
2	Schardl C L， Leuchtmann A， Chung K R， et al. Coevolution by common descent of fungal symbionts （Epichloë spp.） and grass hosts. Molecular， Biology and Evolution， 1997， 14（2）： 133-143.
3	Clay K. Fungal endophytes of grasses： A defensive mutualism between plants and fungi. Ecology， 1998， 69（1）： 10-16.
4	Li C J. Biological and ecological characteristics of Achnatherum inebrians/Neotyphodium endophyte symbiont. Lanzhou： Lanzhou University， 2005.
	李春杰. 醉马草-内生真菌共生体生物学与生态学特性的研究. 兰州：兰州大学， 2005.
5	Li H Q. Effects of endophyte on growth and drought resistance of perennial ryegrass. Lanzhou： Lanzhou University， 2016.
	李会强.内生真菌对多年生黑麦草生长及抗旱性能的影响. 兰州：兰州大学， 2016.
6	Wang Z F， Li C J， White J. Effects of Epichloë endophyte infection on growth， physiological properties and seed germination of wild barley under saline conditions. Journal of Agronomy and Crop Science， 2020， 206（1）： 43-51.
7	Chen Z J， Li C J， Nan Z B， et al. Segregation of Lolium perenne into a subpopulation with high infection by endophyte Epichloëfestucae var. lolii results in improved agronomic performance. Plant and Soil， 2020， 466： 595-612.
8	Chen N， He R L， Chai Q， et al. Transcriptomic analyses giving insights into molecular regulation mechanisms involved in cold tolerance by Epichloë endophyte in seed germination of Achnatherum inebrians.Plant Growth Regulation， 2016， 80（3）： 367-375.
9	Zhang X X. Response of Achnatherum inebrians/Neotyphodium gansuense symbiont to stresses and secondary metabolites activities. Lanzhou： Lanzhou University， 2012.
	张兴旭. 醉马草—内生真菌共生体对胁迫的响应及其次生代谢产物活性的研究. 兰州：兰州大学， 2012.
10	Zhang X X. Effects of endophyte infection on pest resistance to drunken horse grass. Lanzhou： Lanzhou University， 2008.
	张兴旭. 内生真菌对醉马草抗虫性影响的研究. 兰州：兰州大学， 2008.
11	Xia C， Li N， Zhang X X， et al. An Epichloë， endophyte improves photosynthetic ability and dry matter production of its host Achnatherum inebrians， infected by Blumeria graminis， under various soil water conditions. Fungal Ecology， 2016， 22： 26-34.
12	Guo C H， Li X Z， Liu L， et al. Effect of the Epichloë endophyte on the soil nematode community in the rhizosphere of Achnatherum inebrians. Acta Prataculturae Sinica， 2016， 25（4）： 140-148.
	郭长辉，李秀璋，柳莉，等. 内生真菌对醉马草根际土壤线虫群落的影响. 草业学报， 2016， 25（4）： 140-148.
13	Xia C. Responses of Epichloё gansuensis-Achnatherum inebrians symbiont to drought stress. Lanzhou： Lanzhou University， 2018.
	夏超. 醉马草-内生真菌共生体对干旱胁迫的响应. 兰州：兰州大学， 2018.
14	Nan Z B， Li C J. Roles of the grass-Neotyphodium association in pastoral agriculture systems. Acta Ecologica Sinica， 2004， 24（3）： 605-616.
	南志标，李春杰. 禾草-内生真菌共生体在草地农业系统中的作用. 生态学报， 2004， 24（3）： 605-616.
15	Zhang J F. Effect of fungal endophyte on intraspecific competition of Elymus nutans under water stress. Lanzhou： Lanzhou University， 2013.
	张金锋.水分胁迫下内生真菌对垂穗披碱草种内竞争的影响. 兰州：兰州大学， 2013.
16	Bewley J D， Black M. Seed： Seed development and maturation. New York： Plenum Press， 1994.
17	Zhang H S， Hu J. Seed science. Beijing： Science Press， 2015.
	张红生，胡晋. 种子学. 北京：科学出版社， 2015.
18	Ma H Y， Liang Z W. Effects of different soil pH and soil extracts on the germination and seedling growth of Leymus chinensis. Chinese Bulletin of Botany， 2007， 24（2）： 181-188.
	马红媛，梁正伟. 不同pH值土壤及其浸提液对羊草种子萌发和幼苗生长的影响. 植物学通报， 2007， 24（2）： 181-188.
19	Weitbrecht K， Muller K， Leubner M G. First off the mark： Early seed germination. Journal of Experimental Botany， 2011， 62： 3289-3309.
20	Liu H. Study on the resistance and utilization of Medicago sativa L. cv. “Zhongmu No.1”. Lanzhou： Gansu Agricultural University， 2004.
	刘华. 中苜一号紫花苜蓿抗逆性及其利用研究. 兰州：甘肃农业大学， 2004.
21	Saikkonen K， Ahlholm J， Helander M， et al. Endophytic fungi in wild and cultivated grasses in Finland. Ecography， 2000， 23（3）： 360-366.
22	Li C J， Lang M X， Chen Z J， et al. Advances in artificial inoculation technology for grass-endophytic fungi. Acta Prataculturae Sinica， 2021， 30（7）： 179-189.
	李春杰，郎鸣晓，陈振江，等. 禾草-内生真菌人工接种技术研究进展. 草业学报， 2021， 30（7）： 179-189.
23	Philipson M N， Christey M C. The relationship of host and endophyte during flowering seed formation and germination of Lolium perenne. New Zealand Journal of Botany， 1986， 24： 125-134.
24	White J F， Morgen J G， Morrow A C. Taxonomy， life cycle， reproduction and detection of Acremonium endophytes. Agriculture Ecosystems & Environment， 1993， 44（1/2/3/4）： 13-37.
25	Pedroe E G， Pablo H M， Claudio M G. Effects of the Neotyphodium endophyte fungus on dormancy and germination rate of Lolium multiflorum seeds. Austral Ecology， 2006， 31（6）： 767-775.
26	De Battista J P， Bouton J H， Bacon C W， et al. Rhizome and herbage production of endophyte-removed tall fescue clones and populations. Agronomy Journal， 1990， 82（4）： 651-654.
27	Arachevaleta M， Bacon C W， Hoveland C S， et al. Effect of tall fescue endophyte on plant response to environmental stress. Agronomy Journal， 1989， 81（1）： 83-90.
28	Ma B H， Lin W H， Gao M， et al. Effects of salicylic acid and Epichloë on perennial ryegrass （Lolium perenne） under drought stress. Acta Prataculturae Sinica， 2020， 29（1）： 135-144.
	马碧花，蔺伟虎，高敏，等. 干旱胁迫下水杨酸和内生真菌对多年生黑麦草的影响. 草业学报， 2020， 29（1）： 135-144.
29	Malinowski D， Leuchtmann A， Schmidt D， et al. Growth and water status in meadow fescue is affected by Neotyphodium and Phialophora species endophytes. Journal of Agromedicine， 1997， 89： 673-678.
30	Ren A Z， Gao Y B， Gao W S. Effects of endophyte infection on seed germination， seeding growth and osmotic stress resistance of perennial ryegrass （Lolium perenne L.）. Chinese Journal of Plant Ecology， 2002（4）： 420-426.
	任安芝，高玉葆，高文生. 内生真菌侵染对黑麦草种子萌发、幼苗生长及渗透胁迫抗性的影响. 植物生态学报， 2002（4）： 420-426.
31	White J R， Belanger F， Meyer W， et al. Clavicipitalean fungal epibionts and endophytes development of symbiotic interactions with plants. Symbiosis， 2002， 33： 201-213.
32	Clay K. Effects of fungal endophytes on the seed and seedling biology of Lolium perenne and Festuca arundinacea. Oecologia， 1987， 73（3）： 358-362.
33	Zhou J L. Interaction of nitrogen and phosphorus supply and Epichloë bromicola on growth of wild barley. Lanzhou： Lanzhou University， 2019.
	周景乐. 氮磷供应与内生真菌互作对野大麦生长的影响. 兰州：兰州大学， 2019.
34	Chen W， Liu H， Jiang N， et al. Physio-ecological effects of endophyte infection on tall fescue with elevated CO₂ and nitrogen supply. Acta Scientiarum Naturalium Universitatis Nankaiensis， 2015， 48（4）： 43-52.
	陈薇，刘慧，姜楠，等. 不同CO₂浓度和氮素处理下内生真菌感染对高羊茅的生理生态影响. 南开大学学报（自然科学版）， 2015， 48（4）： 43-52.
35	Pang M T. Effect on seed germination and proline metabolism of alfalfa seedlings under PEG stress. Taigu： Shanxi Agricultural University， 2016.
	庞妙甜. PEG胁迫对苜蓿种子萌发及幼苗脯氨酸代谢关键酶活性的影响. 太谷：山西农业大学， 2016.
36	Bu Y Y， Guo P， Ji Y L， et al. Effects of Epichloësinica on Roegneria kamoji seedling physiology under PEG-6000 simulated drought stress. Symbiosis， 2018， 77（2）： 123-132.
37	Li X Z， Simpson W R， Song M L， et al. Effects of seed moisture content and Epichloë endophyte on germination and physiology of Achnatherum inebrians. South African Journal of Botany， 2020， 134（5）： 407-414.
38	Ren A Z. Physiological-ecological mechanisms of endophyte effect on drought adaptation of Lolium perenne L. Tianjing： Nankai University， 2003.
	任安芝. 内生真菌对黑麦草抗旱性影响的生理生态机制研究. 天津：南开大学， 2003.
39	Bacon C W. Abiotic stress tolerances （moisture， nutrients） and photosynthesis in endophyte-infected tall fescue. Agriculture， Ecosystems & Environment， 1993， 44： 123-141.
40	Li C J， Yao X， Nan Z B. Advances in research of Achnatherum inebrians-Epichloë endophyte symboints. Chinese Journal of Plant Ecology， 2018， 42（8）： 793-805.
	李春杰，姚祥，南志标. 醉马草内生真菌共生体研究进展. 植物生态学报， 2018， 42（8）： 793-805.
41	Song M L， Li C J， Peng Q Q， et al. Effects of Neotyphodium endophyte on germination of Hordeum brevisubulatum under temperature and water stress conditions. Acta Agrestia Sinica， 2010， 18（6）： 833-837.
	宋梅玲，李春杰，彭清青，等. 温度和水分胁迫下内生真菌对野大麦种子发芽的影响. 草地学报， 2010， 18（6）： 833-837.
42	Tang K， Zhu W W， Zhou W X， et al. Research progress on effects of soil pH on plant growth and development. Crop Research， 2013， 27（2）： 207-212.
	唐琨，朱伟文，周文新，等. 土壤pH对植物生长发育影响的研究进展. 作物研究， 2013， 27（2）： 207-212.
43	Zhao Y K， Zhang W S， Wang Y N， et al. Research progress in physiology and molecular biology of plant responses to high pH. Chinese Journal of Eco-Agriculture， 2008， 16（3）： 783-787.
	赵彦坤，张文胜，王幼宁，等. 高pH对植物生长发育的影响及其分子生物学研究进展.中国生态农业学报， 2008， 16（3）： 783-787.
44	Peng Q Q， Li C J， Song M L， et al. Effects of Neotyphodium endophytes on seed germination of three grass species under different pH conditions. Acta Prataculturae Sinica， 2011， 20（5）： 72-78.
	彭清青，李春杰，宋梅玲，等. 不同酸碱条件下内生真菌对三种禾草种子萌发的影响. 草业学报， 2011， 20（5）： 72-78.
45	Chen Z J， Liu J， Wei X K， et al. Effects of latosols extracts with different pH and endophytic fungi on growth and physiology of Lolium perenne seedling. Acta Botanica Boreali-Occidentalia Sinica， 2017， 37（7）： 1348-1356.
	陈振江，刘静，魏学凯，等. 不同pH砖红壤浸提液和内生真菌对黑麦草幼苗生长生理的影响. 西北植物学报， 2017， 37（7）： 1348-1356.
46	Wan Z W. Effect of temperature， illumination and pH factor on growth and contents of ergot alkaloids of Epichloë endophyte symbiotic Achnatherum inebrians. Lanzhou： Lanzhou University， 2017.
	万志文. 温度、光照和pH对醉马草内生真菌共生体生长及麦角生物碱含量的影响. 兰州：兰州大学， 2017.
47	Chen Z J， Jin Y Y， Yao X， et al. Fungal endophyte improves survival of Lolium perenne in low fertility soils by increasing root growth， metabolic activity and absorption of nutrients. Plant and Soil， 2020， 452（1/2）： 185-206.
48	Cabot C， Sibole J V， Poschenrieder C. Lessons from crop plants struggling with salinity. Plant Science， 2014， 226（3）： 2-13.
49	Shu K， Qi Y， Chen F， et al. Salt stress represses soybean seed germination by negatively regulating GA biosynthesis while positively mediating ABA biosynthesis. Frontiers in Plant Science， 2017， 8： 1-12.
50	Liu Y W， Yu Y， Fang J. Saline-alkali stress and molecular mechanism of saline-alkali tolerance in plants. Soils and Crops， 2018， 7（2）： 201-211.
	刘奕媺，于洋，方军. 盐碱胁迫及植物耐盐碱分子机制研究. 土壤与作物， 2018， 7（2）： 201-211.
51	Zhang J J， An S Z， Shi C， et al. Effects of endophytic fungi on seed germination and seedling physiological characteristics of Melica transsilvanica under salt stress. Chinese Journal of Grassland， 2017， 39（2）： 59-64.
	张晶晶，安沙舟，施宠，等. 内生真菌侵染对盐胁迫下德兰臭草种子萌发及幼苗生理特性的影响. 中国草地学报， 2017， 39（2）： 59-64.
52	Ahmad R Z， Khalid R， Aqeel M， et al. Fungal endophytes trigger Achnatherum inebrians germination ability against environmental stresses. South African Journal of Botany， 2020（134）： 230-236.
53	Kuang Y， Nan Z B， Tian P. Effects of Epichloë endophyte and seed hydro-priming on the germination of Festuca sinensis under NaCl stress. Acta Prataculturae Sinica， 2016， 25（2）： 160-168.
	旷宇，南志标，田沛. 内生真菌和水引发对NaCl胁迫条件下中华羊茅种子萌发的影响. 草业学报， 2016， 25（2）： 160-168.
54	Zhang P P， Hu L X， Fu J M. Effects of endophytic fungi on seed germination of perennial ryegrass under salt stress. Pratacultural Science， 2012， 29（7）： 1094-1099.
	张萍萍，胡龙兴，傅金民. 内生真菌侵染对盐胁迫下黑麦草种子萌发的影响. 草业科学， 2012， 29（7）： 1094-1099.
55	Zhang Y W. Effect of Epichloë endophyte on seed associated fungi and disease resistance of wild barley under salt stress conditions. Lanzhou： Lanzhou University， 2019.
	张永雯. 内生真菌对野大麦种带真菌及其盐胁迫条件下抗病性的影响. 兰州：兰州大学， 2019.
56	Zhang X X， Nan Z B， Li C J. Research progress of improved resistance of the grass to the heavy metal stress by endophyte. Pratacultural Science， 2014， 31（8）： 1466-1474.
	张兴旭，南志标，李春杰. 内生真菌提高禾草耐重金属胁迫的研究进展. 草业科学， 2014， 31（8）： 1466-1474.
57	Wan Z W， Wang P， Zhang X X， et al. Effect of manganese and SA on germination of Achnatherum inebrians seed containing or free of endophyte. Acta Agrestia Sinica， 2016， 24（1）： 107-113.
	万志文，王萍，张兴旭，等. 锰、水杨酸与内生真菌互作对醉马草种子萌发的影响. 草地学报， 2016， 24（1）： 107-113.
58	Li K， Shi C， Wang W Q， et al. Seed germination and growth effects of endophyte infection on Melica transsilvanica under Pb stress. Journal of Agricultural Resources and Environment， 2020， 37（2）： 280-286.
	李柯，施宠，王文全，等. 重金属Pb胁迫下内生真菌侵染对德兰臭草种子萌发及生长的影响. 农业资源与环境学报， 2020， 37（2）： 280-286.
59	Li K， Shi C， Li H Y， et al. Seed germination and growth effects of endophyte infection on Melica transsilvanica under Cd stress. Xinjiang Agricultural Sciences， 2018， 55（10）： 1916-1926.
	李柯，施宠，李昊宇，等. 重金属Cd胁迫下内生真菌感染对德兰臭草种子萌发及生长的影响. 新疆农业科学， 2018， 55（10）： 1916-1926.
60	Yu H R， Li Q F， He Y M， et al. Effects of different temperature and light on physiological and biochemical characteristics of Clematis fruticosa Turcz. seed germination. Heilongjiang Animal Science and Veterinary Medicine， 2018， 549（9）： 139-142， 146.
	于浩然，李青丰，贺一鸣，等. 不同温度及光照对灌木铁线莲种子萌发生理生化特性的影响. 黑龙江畜牧兽医， 2018， 549（9）： 139-142， 146.
61	Liu K， Baskin J M， Baskin C C， et al. Effect of diurnal fluctuating versus constant temperatures on germination of 445 species from the eastern Tibet Plateau. PLoS One， 2013， 8（7）： e69364.
62	Peng Q Q. Effect of Neotyphodium endophyte on chilling tolerance to Festuca sinensis. Lanzhou： Lanzhou University， 2012.
	彭清青. Neotyphodium内生真菌对中华羊茅耐寒性的影响. 兰州：兰州大学， 2012.
63	Chen N. Molecular mechanism involved in low temperature resistance of endophyte infected drunken horse grass during seed germination. Lanzhou： Lanzhou University， 2011.
	陈娜. 内生真菌提高醉马草低温萌发能力的分子机制. 兰州：兰州大学， 2011.
64	Li X Z， Yao X， Li C J， et al. Potential analysis of grass endophyte Neotyphodium as biocontrol agents. Chinese Journal of Plant Ecology， 2015， 39（6）： 621-634.
	李秀璋，姚祥，李春杰，等. 禾草内生真菌作为生防因子的潜力分析. 植物生态学报， 2015， 39（6）： 621-634.
65	Liu J. Studies on antifungal and insect-resistant activities and fermentation technology of Epichloë grass endophyte. Lanzhou： Lanzhou University， 2019.
	刘静. 禾草内生真菌抑菌抗虫活性和发酵条件的研究. 兰州：兰州大学， 2019.
66	Ren A Z， Gao Y B. Plant endophytic fungi-a kind of resource microorganisms with broad application prospects. Microbiology China， 2001， 28（6）： 90-93.
	任安芝，高玉葆. 植物内生真菌-一类应用前景广阔的资源微生物. 微生物学通报， 2001， 28（6）： 90-93.
67	Li X Z， Song M L， Xiang Y， et al. The effect of seed-borne fungi and Epichloë endophyte on seed germination and biomass of Elymus sibiricus. Frontiers in Microbiology， 2017， 8： 2488.
68	Li C J， Gao J H， Nan Z B. Interactions of Neotyphodium ganauense， Achnatherum inebrians， and plant-pathogenic fungi. Mycological Research， 2007， 111（10）： 1220-1227.
69	Li X Z. Study on the evolution and interactions of Epichloë gansuensis with host seed-borne fungi and rhizospheric microorganism. Lanzhou： Lanzhou University， 2017.
	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. 兰州：兰州大学， 2017.
70	Malinowski D P， Belesky D P. Adaptations of endophyte-infected cool-season grasses to environmental stresses： Mechanisms of drought and mineral stress tolerance. Crop Science， 2000， 40： 923-940.
71	Ma M Z， Christensen M J， Nan Z B. Effects of the endophyte Epichloëfestucae var. lolii of perennial ryegrass （Lolium perenne） on indicators of oxidative stress from pathogenic fungi during seed germination and seedling growth. European Journal of Plant Pathology， 2015， 141（3）： 571-583.
72	Chen F， Meng Y J， Shuai H W， et al. Effect of plant allelochemicals on seed germination and its ecological significance. Chinese Journal of Eco-Agriculture， 2017， 25（1）： 36-46.
	陈锋，孟永杰，帅海威，等. 植物化感物质对种子萌发的影响及其生态学意义. 中国生态农业学报， 2017， 25（1）： 36-46.
73	Sun W H， Yu S W. Allelopathy and its potential application. Plant Physiology Communication， 1992， 28（2）： 81-87.
	孙文浩，余叔文. 相生相克效应及其应用. 植物生理学通讯， 1992， 28（2）： 81-87.
74	Farooq M， Jabran K， Cheema Z A， et al. The role of allelopathy in agricultural pest management. Pest Management Science， 2011， 67（5）： 493-506.
75	Yang S， Li C J， Chai Q， et al. Allelopathic effects of Neotyphodium endophytes of Elymus dahuricus on seeds and seedlings of three turf grass species. Acta Prataculturae Sinica， 2010， 19（4）： 33-40.
	杨松，李春杰，柴青，等. 披碱草内生真菌对三种草坪草种子与种苗的化感效应. 草业学报， 2010， 19（4）： 33-40.
76	Wu L J， Jing Y F， Ren A Z， et al. Effects of endophyte-infected Leymus chinensis on seed germination of Stipa krylovii. Bulletin of Botanical Research， 2012， 32（6）： 669-674.
	吴连杰，荆元芳，任安芝，等. 感染内生真菌的羊草对克氏针茅种子萌发的影响. 植物研究， 2012， 32（6）： 669-674.
77	Yang S， Huang X， Chai Q， et al. Allelopathic effects of endophytic fungi of Achnatherum inebrians Keng on the seed and seedling of three turf grasses. Acta Agrestia Sinica， 2010， 18（1）： 78-83， 88.
	杨松，黄玺，柴青，等. 醉马草内生真菌对3种草坪草种子与种苗的化感效应. 草地学报， 2010， 18（1）： 78-83， 88.
78	Li M M， Gu L J， Ma B H， et al. Effects of Festuca sinensis-endophyte symbiont on germination and seedling growth of Lolium perenne. Grassland and Turf， 2019， 39（1）： 35-42.
	李苗苗，古丽君，马碧花，等. 中华羊茅内生真菌共生体对多年生黑麦草种子萌发和幼苗生长的影响. 草原与草坪， 2019， 39（1）： 35-42.
79	Liu J， Zhou J L， Chen Z J， et al. Effect of seed soaking with Epichloë fermentation broth on germination characteristics in perennial ryegrass under drought stress. Pratacultural Science， 2018， 35（4）： 839-847.
	刘静，周景乐，陈振江，等. 内生真菌发酵液浸种对干旱胁迫下黑麦草种子萌发的影响. 草业科学， 2018， 35（4）： 839-847.
80	Wang Q Y. Effects of fermentation filtrate of Coniothyrium minitans on sunflower and the changes of SOD activity. Journal of Shanxi Agricultural Sciences， 2014， 42（12）： 1274-1275， 1282.
	王庆云. 盾壳霉发酵物对向日葵的促生作用及SOD活性的影响. 山西农业科学， 2014， 42（12）： 1274-1275， 1282.
81	Zhong S， Wang W N， Zhou L Y， et al. Effect of fermentation liquid of Epichloë isolated from Festuca sinensis on seed germination. Journal of Shanxi Agricultural Sciences， 2017， 45（5）： 736-739.
	钟松，王文妮，周连玉，等. 中华羊茅内生真菌发酵液对种子萌发特性的影响.山西农业科学， 2017， 45（5）： 736-739.
82	Xia C， Zhong R， Zhang X X， et al. Allelopathic effects of volatile compounds from endophyte-free and infected Achnatherum inebrians on Lolium perenne. Pratacultural Science， 2015， 32（5）： 658-666.
	夏超，钟睿，张兴旭，等. 醉马草挥发油对多年生黑麦草种子萌发及幼苗生理变化的影响. 草业科学， 2015， 32（5）： 658-666.
83	Huang X， Li C J， Nan Z B， et al. Effects of Achnatherum inebrians infected with Neotyphodium endophyte on accompanying species of Stipa capillata and Poa sphondylodes. Acta Prataculturae Sinica， 2010， 19（5）： 87-93.
	黄玺，李春杰，南志标，等. 醉马草内生真菌对其伴生种硬质早熟禾和针茅生长的影响. 草业学报， 2010， 19（5）： 87-93.
84	Peng S L， Shao H. Research significance and foreground of allelopathy. Chinese Journal of Applied Ecology， 2001， 12（5）： 780-786.
	彭少麟，邵华. 化感作用的研究意义及发展前景. 应用生态学报， 2001， 12（5）： 780-786.
85	Kong C H， Xu T， Hu F S. Study on interactions among allelochemicals of Ageratum conyzoides. Acta Phytoecologica Sinica， 1998， 22（5）： 403-408.
	孔垂华，徐涛，胡飞胜. 胜红蓟化感物质之间相互作用的研究. 植物生态学报， 1998， 22（5）： 403-408.
86	Garbutt K， Williams W E， Bazzaz F A. Analysis of the differential response of five annuals to elevated CO₂ during growth. Ecology， 1990， 71（3）： 1185-1194.
87	Shi Z B， Zhou Y， Li X， et al. Physio-ecological effects of endophyte infection on the host grass with elevated CO₂. Acta Ecologica Sinica， 2013， 33（19）： 6135-6141.
	师志冰，周勇，李夏，等. CO₂浓度升高条件下内生真菌感染对宿主植物的生理生态影响. 生态学报， 2013， 33（19）： 6135-6141.
88	Zhang X X， Li C J， Nan Z B. Interactive effect of storage time and endophytic fungi on seed physiology of Achnatherum inebrians （Hance） Ken. Acta Agrestia Sinica， 2010， 18（2）： 252-257.
	张兴旭，李春杰，南志标. 内生真菌和保存时间互作对醉马草种子的生理影响. 草地学报， 2010， 18（2）： 252-257.
89	Jin W J， Li C J. Effect of storage time on the activity of endophytic fungi of Festuca sinensis. Chinese Journal of Biological Control， 2015， 31（6）： 930-935.
	金文进，李春杰. 保存时间对中华羊茅种子内生真菌活力的影响.中国生物防治学报， 2015， 31（6）： 930-935.
90	Hume D E， Card S D， Bylin A G， et al. Endophyte storage and seed germination of Epichloe-infected meadow fescue. Seed Science & Technology， 2016， 44（1）： 138-155.
91	Li J L. Research progress in physiology and ecology of turfgrass resistance. Nanjing： Nanjing University Press， 2017.
	李建龙. 草坪草抗性生理与生态研究进展. 南京：南京大学出版社， 2017.
92	Shi Y. Effects of endophytic fungi on growth and development of plants. Modern Agricultural Science and Technology， 2010（6）： 36-38.
	石瑛. 内生真菌对植物生长发育的影响. 现代农业科技， 2010（6）： 36-38.
93	Dai M L， Wang P， Sun J K， et al. Research progress of saline-alkali stress effect on seeds’ germination and its physiological and biochemical mechanism. Northern Horticulture， 2015（10）： 176-179.
	代明龙，王平，孙吉康，等. 盐碱胁迫对植物种子萌发的影响及生理生化机制研究进展. 北方园艺， 2015（10）： 176-179.
94	Cai M F， Li K M， Xie D P， et al. The status and protection strategy of farmland soils polluted by heavy metals. Environmental Science & Technology， 2014， 37（120）： 223-230.
	蔡美芳，李开明，谢丹平，等. 我国耕地土壤重金属污染现状与防治对策研究. 环境科学与技术， 2014， 37（120）： 223-230.
95	Nancy V， Aleta M. Testing native species with deep dormancy. Seed and Technology， 2002， 24（1）： 43-51.
96	Li C J， Wang Z F， Chen T X， et al. Creation of novel barley germplasm using an Epichloë endophyte. Chinese Science Bulletin， 2021， 66（20）： 2608-2617.
	李春杰，王正凤，陈泰祥，等. 利用禾草内生真菌创制大麦新种质. 科学通报， 2021， 66（20）： 2608-2617.
97	Chen L. Molecular detection， genotypes and chemotypes of Epichloë endophytes in Achnatherum inebrians. Lanzhou： Lanzhou University， 2017.
	陈丽. 醉马草内生真菌分子检测、基因型及产碱多样性的研究. 兰州：兰州大学， 2015.

胁迫 Stress	宿主 Host	对宿主种子萌发的影响 Effect on seed germination of hosts	参考文献 Reference	备注 Remarks
渗透 Osmosis	黑麦草L. perenne	E+相较于E-发芽率提高、发芽时间缩短。E+ has higher germination rate and shorter germination time than E-.	［30］
	鹅观草R. kamoji	E+相较于E-发芽率提高、发芽时间缩短。E+ has higher germination rate and shorter germination time than E-.	［36］
	醉马草A. inebrians	E+发芽率和发芽指数高于E-。E+ germination rate and germination index are higher than E-.	［37］
	高羊茅F. arundinacea	E+发芽率低于E-。E+ germination rate is lower than E-.	［39］
酸碱 pH	醉马草A.inebrians	E+发芽率、发芽势高于E-。E+ germination rate and germination potential are higher than E-.	［44］
	野大麦H. brevisubulatum	E+发芽势显著高于E-。E+ germination potential is significantly higher than E-.	［44］
	中华羊茅F.sinensis	E+发芽率、发芽势高于E-。E+ germination rate and germination potential are higher than E-.	［44］
	黑麦草L. perenne	E+发芽率下降趋势更缓。E+ germination rate decreased more slowly.	［45］
盐分 Salt	野大麦H. brevisubulatum	E+发芽率、发芽指数高于E-。E+ germination rate and germination index are higher than E-.	［6］
	德兰臭草M. transsilvanica	E+与E-之间发芽率、相对萌发率与萌发指数差异显著。There are significant differences in germination rate， relative germination rate and germination index between E+ and E-.	［51］
	醉马草A. inebrians	E+发芽率、发芽指数高于E-。E+ germination rate and germination index are higher than E-.	［52］
	中华羊茅F.sinensis	E+萌发力及幼苗生长高于E-。E+ germination power and seedling growth are higher than E-.	［53］
	黑麦草L. perenne	E+萌发率、活力指数、苗长高于E-。E+ germination rate， vitality index and seedling length are higher than E-.	［54］
重金属 Heavy metal	醉马草A.inebrians	E+种子对锰耐受力大于E-。E+ seeds are more resistant to Mn²⁺ than E-.	［57］	锰Mn
	醉马草A.inebrians	E+萌发率、发芽指数高于E-。E+ germination rate and germination index are higher than E-.	［9］	镉Cd
	德兰臭草M.transsilvanica	在高Pb浓度下，E+种子各项萌发指标高于E-。E+ germination parameters are higher than E- under high Pb concentration.	［58］	铅Pb
	德兰臭草M.transsilvanica	E+发芽率、发芽势、活力指数高于E-。E+ germination rate， germination potential and vitality index are higher than E-.	［59］	镉Cd
	披碱草E. dahuricus	E+萌发率高于E-。E+ germination rate is higher than E-.	［56］	镉Cd
温度 Temperature	野大麦H.brevisubulatum	高温（30 ℃）和低温（10 ℃）胁迫下，E+发芽率、发芽指数高于E-。E+ germination rate and germination index are higher than E- under high temperature （30 ℃） and low temperature （10 ℃） stress.	［41］
	中华羊茅F.sinensis	5~30 ℃条件下E+发芽率、幼苗干重、胚芽生长高于E-，萌发时间小于E-。E+ germination rate， seedling dry weight， embryo growth are higher than E-， seed germination time is shorter than E- under 5-30 ℃.	［62］
	醉马草A.inebrians	10 ℃下E+发芽率高于E-。E+ germination rate is higher than E- under 10 ℃.	［63］
生物 Biology	野大麦H.brevisubulatum	E+种子萌发指标高于E-。E+ germination parameters are higher than E-.	［55］	#
生物 Biology	黑麦草L. perenne	E+萌发率、发芽指数高于E-。E+ germination rate and germination index are higher than E-.	［71］	##

胁迫 Stress	宿主 Host	对宿主种子萌发的影响 Effect on seed germination of hosts	参考文献 Reference	备注 Remarks
渗透 Osmosis	黑麦草L. perenne	E+相较于E-发芽率提高、发芽时间缩短。E+ has higher germination rate and shorter germination time than E-.	［30］
	鹅观草R. kamoji	E+相较于E-发芽率提高、发芽时间缩短。E+ has higher germination rate and shorter germination time than E-.	［36］
	醉马草A. inebrians	E+发芽率和发芽指数高于E-。E+ germination rate and germination index are higher than E-.	［37］
	高羊茅F. arundinacea	E+发芽率低于E-。E+ germination rate is lower than E-.	［39］
酸碱 pH	醉马草A.inebrians	E+发芽率、发芽势高于E-。E+ germination rate and germination potential are higher than E-.	［44］
	野大麦H. brevisubulatum	E+发芽势显著高于E-。E+ germination potential is significantly higher than E-.	［44］
	中华羊茅F.sinensis	E+发芽率、发芽势高于E-。E+ germination rate and germination potential are higher than E-.	［44］
	黑麦草L. perenne	E+发芽率下降趋势更缓。E+ germination rate decreased more slowly.	［45］
盐分 Salt	野大麦H. brevisubulatum	E+发芽率、发芽指数高于E-。E+ germination rate and germination index are higher than E-.	［6］
	德兰臭草M. transsilvanica	E+与E-之间发芽率、相对萌发率与萌发指数差异显著。There are significant differences in germination rate， relative germination rate and germination index between E+ and E-.	［51］
	醉马草A. inebrians	E+发芽率、发芽指数高于E-。E+ germination rate and germination index are higher than E-.	［52］
	中华羊茅F.sinensis	E+萌发力及幼苗生长高于E-。E+ germination power and seedling growth are higher than E-.	［53］
	黑麦草L. perenne	E+萌发率、活力指数、苗长高于E-。E+ germination rate， vitality index and seedling length are higher than E-.	［54］
重金属 Heavy metal	醉马草A.inebrians	E+种子对锰耐受力大于E-。E+ seeds are more resistant to Mn²⁺ than E-.	［57］	锰Mn
	醉马草A.inebrians	E+萌发率、发芽指数高于E-。E+ germination rate and germination index are higher than E-.	［9］	镉Cd
	德兰臭草M.transsilvanica	在高Pb浓度下，E+种子各项萌发指标高于E-。E+ germination parameters are higher than E- under high Pb concentration.	［58］	铅Pb
	德兰臭草M.transsilvanica	E+发芽率、发芽势、活力指数高于E-。E+ germination rate， germination potential and vitality index are higher than E-.	［59］	镉Cd
	披碱草E. dahuricus	E+萌发率高于E-。E+ germination rate is higher than E-.	［56］	镉Cd
温度 Temperature	野大麦H.brevisubulatum	高温（30 ℃）和低温（10 ℃）胁迫下，E+发芽率、发芽指数高于E-。E+ germination rate and germination index are higher than E- under high temperature （30 ℃） and low temperature （10 ℃） stress.	［41］
	中华羊茅F.sinensis	5~30 ℃条件下E+发芽率、幼苗干重、胚芽生长高于E-，萌发时间小于E-。E+ germination rate， seedling dry weight， embryo growth are higher than E-， seed germination time is shorter than E- under 5-30 ℃.	［62］
	醉马草A.inebrians	10 ℃下E+发芽率高于E-。E+ germination rate is higher than E- under 10 ℃.	［63］
生物 Biology	野大麦H.brevisubulatum	E+种子萌发指标高于E-。E+ germination parameters are higher than E-.	［55］	#
生物 Biology	黑麦草L. perenne	E+萌发率、发芽指数高于E-。E+ germination rate and germination index are higher than E-.	［71］	##

[1]	Xin-tong ZHAO, Xiao-dong CHEN, Zi-ji LI, Ju-ming ZHANG, Tian-zeng LIU. An evaluation of the effects of the plant endophyte Enterobacter on the salt tolerance of bermudagrass [J]. Acta Prataculturae Sinica, 2021, 30(9): 127-136.
[2]	Sai Hen-na BAO, Xiang-tao WANG, Jun-xi WU, Yan-jun MIAO, Xiang JIA, Yan-ting TIAN. Allelopathic effect of root extracts from Astragalus strictus seedlings on eight wild plants of Tibet [J]. Acta Prataculturae Sinica, 2021, 30(5): 211-220.
[3]	Li-rong TONG, Shun-gang NI, Ya-nan ZHOU, Juan WANG, Fang-shan XIA. Effects of different seed coating formulations on seed germination and seedling growth of Lespedeza davurica [J]. Acta Prataculturae Sinica, 2021, 30(2): 124-134.
[4]	Min ZHANG, Kanjana NIPAPAN, Ru LI, Yang FU, Dong-sheng TANG. Environmental factors affecting seed germination of prairie grass （Bromus willdenowii） in Yunnan Province [J]. Acta Prataculturae Sinica, 2021, 30(12): 143-151.
[5]	Yu-lian GAO, Jing CHANG, Yi-hui WANG, Feng LI, Hai-ping LI, Chong-yong MA. Allelopathic effects of Stellera chamaejasme on seed germination and growth of three crops [J]. Acta Prataculturae Sinica, 2021, 30(10): 83-91.
[6]	NIU Huan-huan, WANG Sen-sen, JIA Hong-ding, CHEN Gui-hua. Allelopathic effects of extracts of Vicia villosa on the germination of four forage seeds [J]. Acta Prataculturae Sinica, 2020, 29(9): 161-168.
[7]	LI Feng-lan, WU Jia-wen, YAO Shu-kuan, ZHAO Zi-yi, ZHAO Xiao-can, HE Fu-meng, ZHU Yuan-fang, SHI Qi-hai, ZHOU Lei, XU Yong-qing. A study of the allelopathic effect of extracts from different parts of Iva xanthiifolia on five native species [J]. Acta Prataculturae Sinica, 2020, 29(9): 169-178.
[8]	CUI Xue-lian, XIA Chao. Effect of exogenous abscisic acid on seedling establishment of Epichloё gansuensis-Achnatherum inebrians symbiont [J]. Acta Prataculturae Sinica, 2020, 29(7): 70-80.
[9]	LIANG Jun, QUAN Xiao-long, ZHANG Jie-xue, SHI Hui-lan, DUAN Zhong-hua, QIAO You-ming. Potential allelopathic effects of water extracts of three grasses on germination of their own seeds and seedling growth [J]. Acta Prataculturae Sinica, 2020, 29(7): 81-89.
[10]	WANG Ju-hong, SHI Sheng-jing, CHEN Wen, LI Yun, CUI Xian-liang. Allelopathy and invasiveness of Bidens pilosa and Mimosa pudica [J]. Acta Prataculturae Sinica, 2020, 29(4): 81-91.
[11]	ZHANG Jin-yu, ZHOU Zhong-ze, YE Xiao-xin. Allelopathic potential of Carex thunbergii against three Poaceae species [J]. Acta Prataculturae Sinica, 2020, 29(3): 103-111.
[12]	Ju-hong WANG, Sheng-jing SHI, Wen CHEN, Gui-mei GAN, Sai-na CHEN, Zhang-wei LI. Effects of Bacillus subtilis and three actinomycetes on seed germination and seedling growth of Bidens pilosa and Eclipta prostrata under salt stress [J]. Acta Prataculturae Sinica, 2020, 29(12): 112-120.
[13]	WANG Yu-xuan, TANG Zong-shou, CAO Meng-lin, LI Ya-xin, ZHANG Tian-bao, DU Hui-ling. Allelopathic effects of Hylotelephium erythrostictum flowers on three weed species and identification of potential allelochemicals [J]. Acta Prataculturae Sinica, 2020, 29(1): 175-182.
[14]	LI Zhen, YUN Lan, SHI Zi-ying, WANG Jun, ZHANG Chen, GUO Hong-yu, SHENG Yu. Physiological characteristics of Psathyrostachys juncea at seed germination and seedling growth stages under salt stress [J]. Acta Prataculturae Sinica, 2019, 28(8): 119-129.
[15]	LIU Ya-jing, MENG Zhong-ju, DANG Xiao-hong, SONG Wen-juan, ZHAI Bo. Allelopathic effects of Stellera chamaejasme on seed germination and seedling growth of alfalfa and two forage grasses [J]. Acta Prataculturae Sinica, 2019, 28(8): 130-138.