Advances in research of Achnatherum inebrians-Epichloë endophyte symbionts

doi:10.17521/cjpe.2018.0001

Abstract

Abstract:

Research on drunken horse grass (Achnatherum inebrians)-endophytic fungi (Epichloë gansuensis, E. inebrians) represented an important area of endophyte research in China. Compared with tall fescue (Festuca arundinacea)-endophyte (E. coenophiala) in the United States and perennial ryegrass (Lolium perenne)-endophyte (E. festucae var. lolii) in New Zealand, A. inebrians-endophyte research in China is becoming a leading branch of endophyte research. The studies of A. inebrians-endophyte symbionts over the past three decades were reviewed, mainly including endophyte distribution, detection methods, infection rate, fungal diversity, stress tolerances of drought, cold, salt, heavy metal, pest, disease and their mechanisms, secondary metabolite of alkaloids, feeding experiments and its ecological functions etc. Research reveals that drunken horse grass itself has no toxicity, the grazed animals will be intoxicated if the grass bears infection by Epichloë endophytes and produces ergot alkaloids. Future research will focus on E. gansuensis and E. inebrians genomics and endophyte functions. In other research we will create new grass cultivars that are 1) sanitized of the endophyte and non-toxic, 2) utilizing novel endophytes that deter birds feeding and 3) utilizing endophytes that are modified to produce novel fungicides, pesticides or medicines.

Key words: grasses, fungal endophytes, future perspectives, toxic grass control and utilization, bio-control

LI Chun-Jie, YAO Xiang, NAN Zhi-Biao. Advances in research of Achnatherum inebrians-Epichloë endophyte symbionts[J]. Chin J Plan Ecolo, 2018, 42(8): 793-805.

Add to citation manager EndNote|Ris|BibTeX

URL: https://www.plant-ecology.com/EN/10.17521/cjpe.2018.0001

https://www.plant-ecology.com/EN/Y2018/V42/I8/793

Figures/Tables 1

References 109

[1]	Adams AE, Kazenel MR, Rudgers JA ( 2017). Does a foliar endophyte improve plant fitness under flooding? Plant Ecology, 218, 711-723. DOI URL
[2]	Bruehl GW, Kaiser WJ, Klein RE ( 1994). An endophyte of Achnatherum inebrians, an intoxicating grass of northwest China. Mycologia, 86, 773-776.
[3]	Cao GR, Dang XP, Duan DX, Li SJ, Zhou JH ( 1991). Poisoning test of Achnatherum inebrians. Chinese Journal of Veterinary Science and Technology, 21(9), 26-27.
	[ 曹光荣, 党晓鹏, 段得贤, 李绍君, 周进海 ( 1991). 醉马草中毒试验. 中国兽医科技, 21(9), 26-27.]
[4]	Card SD, Rolston MP, Park Z, Cox N, Hume DE ( 2013). Assessment of fluorescein-based fluorescent dyes for tracing Neotyphodium endophytes in plant. Mycologia, 105, 221-229. DOI URL PMID
[5]	Chen L ( 2015). Molecular Detection, Genotypes and Chemotypes of Epichloë endophytes in Achnatherum inebrians. PhD dissertation, Lanzhou University, Lanzhou.
	[ 陈丽 ( 2015). 醉马草内生真菌分子检测、基因型及产碱多样性的研究. 博士学位论文, 兰州大学, 兰州.]
[6]	Chen L, Li XZ, Li CJ, Swoboda GA, Young CA, Sugawara K, Leuchtmann A, Schardl CL ( 2015). Two distinct Epichloë species symbiotic with Achnatherum inebrians, drunken horse grass. Mycologia, 107, 863-873.
[7]	Chen N ( 2008). Genetic Diversity of Drunken Horse Grass (Achnatherum inebrians) and Effects of Its Endophyte Infection on Cold Tolerance. Master degree dissertation, Lanzhou University, Lanzhou.
	[ 陈娜 ( 2008). 醉马草遗传多样性及内生真菌对其抗寒性影响. 硕士学位论文, 兰州大学, 兰州.]
[8]	Chen N ( 2011). Molecular Mechanism Involved in Low Temperature Resistance of Endophyte Infected Drunken Horse Grass During Seed Germination. PhD dissertation, Lanzhou University, Lanzhou.
	[ 陈娜 ( 2011). 内生真菌提高醉马草低温萌发能力的分子机制. 博士学位论文, 兰州大学, 兰州.]
[9]	Chen N, He RL, Chai Q, Li CJ, Nan ZB ( 2016). Transcriptomic analyses giving insights into molecular regulation mechanisms involved in cold tolerance by Epichloë endophyte in seed germination of Achnatherum inebrians. Plant Growth Regulation, 80, 367-375. DOI URL
[10]	Clark EM, White JF, Patterson RM ( 1983). Improved histochemical techniques for the detection of Acremonium coenophialum in tall fescue and methods of in vitro culture of the fungus. Journal of Microbiological Methods, 1, 149-155.
[11]	Clayton W, Eaton CJ, Dupont PY, Gillanders T, Cameron N, Saikia S, Scott B ( 2017). Analysis of simple sequence repeat (SSR) structure and sequence within Epichloë endophyte genomes reveals impacts on gene structure and insights into ancestral hybridization events. PLOS ONE, 12, e0183748. DOI: 10.1371/journal.pone.0183748.
[12]	Dai LY ( 2010). Ergot Alkloids in Symbiont of Achnatherum inebrians and Neotyphodium gansuense. Master degree dissertation, Lanzhou University, Lanzhou.
	[ 代乐英 ( 2010). 醉马草内生真菌共生体麦角生物碱的研究. 硕士学位论文, 兰州大学, 兰州.]
[13]	Dang XP, Cao GR, Duan DX, Li SJ, Zhao XW, Zhou JH ( 1992). Studies on the toxic constituent of Achnatherum inebrians. Acta Veterinaria et Zootechnica Sinica, 23, 366-371.
	[ 党晓鹏, 曹光荣, 段得贤, 李绍君, 赵效文, 周进海 ( 1992). 醉马草的有毒成分研究. 畜牧兽医学报, 23, 366-371.]
[14]	Deng KD, Peng HH, Li WR, Warren BE, Fletcher IG ( 1998). The ergonovine content and the nutritive value of ammoniated drunken horse grass with urea. Pratacultural Science, 15(4), 10-13.
	[ 邓凯东, 彭海宏, 李文蓉, Warren BE, Fletcher IG ( 1998). 尿素氨化醉马草的麦角新碱含量及其营养价值. 草业科学, 15(4), 10-13.]
[15]	Dong LL, An SZ, Jin GL, Xun QL, Wei P, Qu HM ( 2014). Dynamic population changes of Achnatherum inebrians seedlings. Pratacultural Science, 31, 499-503.
	[ 董莉莉, 安沙舟, 靳瑰丽, 荀其蕾, 魏鹏, 屈慧敏 ( 2014). 不同生境下醉马草幼苗数量的变化规律. 草业科学, 31, 499-503.]
[16]	Gao JH ( 2006). Detection of Ergot Alkaloid Production by Neotyphodium gansuense in vitro and Sequencing the dmaW Gene of N.lolii Isolates AR1 and Lp19. Master degree dissertation, Lanzhou University, Lanzhou.
	[ 高嘉卉 ( 2006). Neotyphodium gansuense离体麦角碱的检测和N. lolii麦角碱基因的测序. 硕士学位论文, 兰州大学, 兰州.]
[17]	Gao Y ( 2001). Significant control of glyphosate on Achnatherum inebrians. Chinese Journal of Animal Husbandry and Veterinary Medicine, 5, 8.
	[ 高原 ( 2001). 用草甘膦杀灭醉马草效果明显. 畜牧兽医科技信息, 5, 8.]
[18]	Gou XY ( 2007). Effects of Neotyphodium Endophyte on Salt Tolerance to Drunken Horse Grass (Achnatherum inebrians). Master degree dissertation, Lanzhou University, Lanzhou.
	[ 缑小媛 ( 2007). 内生真菌对醉马草耐盐性的影响研究. 硕士学位论文, 兰州大学, 兰州.]
[19]	Guo CH, Li XZ, Liu L, Cao JX, Li CJ ( 2016). Effect of the Epichloë endophyte on the soil nematode community in the rhizosphere of Achnatherum inebrians. Acta Prataculturae Sinica, 25, 140-148. DOI URL
	[ 郭长辉, 李秀璋, 柳莉, 曹建新, 李春杰 ( 2016). 内生真菌对醉马草根际土壤线虫群落的影响. 草业学报, 25, 140-148.] DOI URL
[20]	Hance HF ( 1876). On a Mongolian grass producing intoxicating in cattle. Journal of Botany, 14, 210-212.
[21]	He MH ( 2001). Investigation on poisoning of sheep. Chinese Qinghai Journal of Animal and Veterinary Science, 31, 27.
	[ 何明海 ( 2001). 放牧绵羊醉马草中毒调查. 青海畜牧兽医杂志, 31, 27.]
[22]	Hu CX ( 2013). Effects of N, P and Tryptophan on Growth and Ergot Alkloid Content in Achnatherum inebrians Symbiotic with Neotyphodium gansuense. Master degree dissertation, Lanzhou University, Lanzhou.
	[ 胡春霞 ( 2013). 氮、磷和色氨酸对醉马草内生真菌共生体生长及麦角生物碱含量的影响. 硕士学位论文, 兰州大学, 兰州.]
[23]	Huang X ( 2011). Effects of Neotyphodium gansuense on Competition of Achnatherum inebrians. Master degree dissertation, Lanzhou University, Lanzhou.
	[ 黄玺 ( 2011). 甘肃内生真菌对醉马草竞争性的影响. 硕士学位论文, 兰州大学, 兰州.]
[24]	Huang X, Li CJ, Nan ZB ( 2012). Competition effects between Medicago sativa and Achnatherum inebrians. Acta Prataculturae Sinica, 21(1), 59-65. DOI URL
	[ 黄玺, 李春杰, 南志标 ( 2012). 紫花苜蓿与醉马草的竞争效应. 草业学报, 21(1), 59-65.] DOI URL
[25]	Huang X, Li CJ, Nan ZB, Yang S, Chai Q ( 2010). Effects of Achnatherum inebrians infected with Neotyphodium endophyte on accompanying species of stipa capillata and Poa sphondylodes. Acta Prataculturae Sinica, 19(5), 87-93. DOI URL
	[ 黄玺, 李春杰, 南志标, 杨松, 柴青 ( 2010). 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, 19(5), 87-93.] DOI URL
[26]	Huang X, Li XZ, Chai Q, Li CJ ( 2013). Effects of Achnatherum inebrians/Neotyphodium endophyte symbiont on microflora and nutrient of soil. Pratacultural Science, 30, 352-356.
	[ 黄玺, 李秀璋, 柴青, 李春杰 ( 2013). 醉马草内生真菌共生体对土壤微生物和养分的影响. 草业科学, 30, 352-356.]
[27]	Jianati, Sahedula HM, Nuerlan ( 1998). Study of sheep feed by Achnatherum inebrians. Journal of Animal Industry in Xinjiang, ( 4), 31.
	[ 贾纳提, 萨赫都拉·霍曼, 努尔兰 ( 1998). 醉马草饲喂试验研究. 新疆畜牧业, ( 4), 31.]
[28]	Jin GL, Dong LL, An SZ, He L, Liang N, Zhang MN ( 2014). Interspecific relationship of Achnatherum inebrians and communities in the north slope of Tianshan Mountain. Acta Agrestia Sinica, 22, 1179-1185. DOI URL
	[ 靳瑰丽, 董莉莉, 安沙舟, 何龙, 梁娜, 张梦妮 ( 2014). 天山北坡醉马草群落种间关联分析. 草地学报, 22, 1179-1185.] DOI URL
[29]	Jin WJ ( 2009). Diversity of Neotyphodium Endophytes Symbiotic with Achnatherum inebrians. Master degree dissertation, Lanzhou University, Lanzhou.
	[ 金文进 ( 2009). 醉马草内生真菌多样性的研究. 硕士学位论文, 兰州大学, 兰州.]
[30]	Jin WJ, Li CJ, Wang ZF ( 2015). Research adcances on diversityof grass Epichloë endophytes. Acta Prataculturae Sinica, 24, 168-175.
	[ 金文进, 李春杰, 王正凤 ( 2015). 内生真菌多样性及其意义. 草业学报, 24, 168-175.]
[31]	Johnson LJ, de Bonth ACM, Briggs LR, Caradus JR, Finch SC, Fleetwood DJ, Fletcher LR, Hume DE, Johnson RD, Popay AJ, Tapper BA, Simpson WR, Voisey CR, Card SD ( 2013). The exploitation of epichloae endophytes for agricultural benefit. Fungal Diversity, 60, 171-188. DOI URL
[32]	Keng YL ( 1959). Illustration of China Main Plants—Poaceae. Science Press, Beijing.
	[ 耿以礼 ( 1959). 中国主要植物图说——禾本科. 科学出版社, 北京.]
[33]	Leuchtmann A, Bacon CW, Schardl CL, White JF, Tadych M ( 2014). Nomenclatural realignment of Neotyphodium species with genus Epichloë. Mycologia, 106, 202-215.
[34]	Li BJ, Zheng XH, Sahedula, Sun SC ( 1996). An investigation of endophyte-grasses in north-west of China. Grassland of China, ( 2), 29-32.
	[ 李保军, 郑晓红, 萨赫都拉, 孙穗长 ( 1996). 新疆部分禾草的植物内生菌调查. 中国草地, ( 2), 29-32.]
[35]	Li CJ ( 2005). Biological and Ecological Characteristics of Achnatherum inebrians/Neotyphodium Endophyte Symbiont. PhD dissertation, Lanzhou University, Lanzhou.
	[ 李春杰 ( 2005). 醉马草-内生真菌共生体生物学与生态学特性的研究. 博士学位论文, 兰州大学, 兰州.]
[36]	Li CJ, Gao JH, Nan ZB ( 2007). Interactions of Neotyphodium gansuense, Achnatherum inebrians and plant-pathogenic fungi. Mycological Research, 111, 1220-1227.
[37]	Li CJ, Nan ZB ( 2009). Detection technology of endophytic fungus in hard grass: China, ZL200410025944.0. 2009-07-08.
	[ 李春杰, 南志标 ( 2009). 硬质禾草内生真菌检测技术: 中国, ZL200410025944.0. 2009-07-08.]
[38]	Li CJ, Nan ZB, Li F ( 2008). Biological and physiological characteristics of Neotyphodium gansuense symbiotic with Achnatherum inebrians. Microbiological Research, 163, 431-440.
[39]	Li CJ, Nan ZB, Liu Y, Volk HP, Dapprich P ( 2008). Methodology of endophyte detection of drunken horse grass (Achnatherum inebrians). Edible Fungi of China, 27, 16-19.
	[ 李春杰, 南志标, 刘勇, VolkH P, Dapprich P ( 2008). 醉马草内生真菌检测方法的研究. 中国食用菌, 27, 16-19.]
[40]	Li CJ, Nan ZB, Volker HP, Dapprich PD, Liu Y ( 2004). A new Neotyphodium species symbiotic with drunken horse grass(Achnatherum inebrians) in China. Mycotaxon, 90, 141-147.
[41]	Li CJ, Nan ZB, Zhang CJ, Zhang CY, Zhang YH ( 2009). Effects of drunken horse grass infected with endophyte on Chinese rabbit. Journal of Agricultural Science and Technology, 11(2), 84-90. DOI URL
	[ 李春杰, 南志标, 张昌吉, 张崇岳, 张燕慧 ( 2009). 醉马草内生真菌对家兔的影响. 中国农业科技导报, 11(2), 84-90.] DOI URL
[42]	Li F ( 2007). Effects of Endophyte Infection on Drought Resistance to Drunken Horse Grass (Achnatherum inebrians). Master degree dissertation, Lanzhou University, Lanzhou.
	[ 李飞 ( 2007). 内生真菌对醉马草抗旱性影响的研究. 硕士学位论文, 兰州大学, 兰州.]
[43]	Li H, Sun H, He Z ( 2015). Achnatherum inebrians straw as a potential raw material for pulp and paper production. Journal of Cleaner Production, 101, 193-196.
[44]	Li NN, Zhao YF, Xia C, Zhong R, Zhang XX ( 2016). Effects of thiophanate methyl on seed borne Epichloë fungal endophyte of Achnatherum inebrians. Pratacultural Science, 33, 1306-1314.
	[ 李娜娜, 赵玉凤, 夏超, 钟睿, 张兴旭 ( 2016). 甲基托布津对醉马草种带内生真菌的灭菌活性. 草业科学, 33, 1306-1314.]
[45]	Li NN, Xia C, Zhong R, Ju YW, Nan ZB, Christensen MJ, Zhang XX ( 2018). Interactive effects of water stress and powdery mildew (Blumeria graminis) on the alkaloid production of Achnatherum inebrians, infected by Epichloë, endophyte. Science China Life Sciences, 61, 1-3.
[46]	Li XS, Ren JZ, Feng KM, Lei TS, Ar YD, Zhang XZ, Muhetaer, Shuanghuer ( 1996). Study on ecological control of Achnatherum inebrians. Acta Prataculturae Sinica, 6(2), 14-17.
	[ 李学森, 任继生, 冯克明, 雷特生, 阿依丁, 张学洲, 穆合塔尔, 双湖尔 ( 1996). 醉马草生态控制的研究. 草业学报, 6(2), 14-17.]
[47]	Li XS, Zhang XZ, Gu X, Zheng Y ( 1998). Relationship of toxic substance in Achnatherum inebrians and endophytic fungi. Grass-feeding Livestock, ( 4), 44-47.
	[ 李学森, 张学洲, 顾祥, 郑瑛 ( 1998). 醉马草有毒物质与其内生真菌的关系. 草食家畜, ( 4), 44-47.]
[48]	Li XZ ( 2017). Study on the Evolution and Interactions of Epichloë gansuensis with Host Seed-borne Fungi and Rhizospheric Microorganism. PhD dissertation, Lanzhou University, Lanzhou.
	[ 李秀璋 ( 2017). 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. 博士学位论文, 兰州大学, 兰州.]
[49]	Liang Y, Wang H, Li C, Nan Z, Li F ( 2017). Effects of feeding drunken horse grass infected with Epichloë gansuensis, endophyte on animal performance, clinical symptoms and physiological parameters in sheep. BMC Veterinary Research, 13, 223. DOI: 10.1186/s12917-017-1120-6. DOI URL PMID
[50]	Matthew AC, Brian AT, Wayne RS, Richard DJ, Wade M, Arvina R, Yonathan L, Pierre-Yves D, Linda JJ, Barry S, Austen RDG, Murray PC ( 2017). Epichloë hybrida, sp. nov., an emerging model system for investigating fungal allopolyploidy. Mycologia, 109, 715-729. DOI URL
[51]	Miles CO, Lane GA, di Menna ME, Garthwaite I, Piper EL, Ball OJP, Latch GCM, Allen JM, Hunt MB, Bush LP, Min FK, Fletcher I, Harris PS ( 1996). High levels of ergonovine and lysergic acid amide in toxic Achnatherum inebrians accompany infection by an Acremonium-like endophytic fungus. Journal of Agricultural and Food Chemistry, 44, 1285-1290.
[52]	Moon CD, Guillaumin JJ, Ravel C, Li C, Craven KD, Schardl CL ( 2007). New Neotyphodium endophyte species from the grass tribes Stipeae and Meliceae. Mycologia, 99, 895-905. DOI URL PMID
[53]	Nan ZB ( 1996 a). Incidence and distribution of endophytic fungi in seeds of some native and introduced grasses in China. Acta Prataculturae Sinica, 5(2), 1-8.
	[ 南志标 ( 1996 a). 内生真菌在我国部分国产和引进禾草品种种子中的分布. 草业学报, 5(2), 1-8.]
[54]	Nan ZB ( 1996 b). Incidence and distribution of endophytic fungi in seedlings and plants of some native and introduced grasses in China. Acta Prataculturae Sinica, 5(3), 13-17.
	[ 南志标 ( 1996 b). 内生真菌在我国部分国产和引进禾草品种的幼苗及成株中的分布. 草业学报, 5(3), 13-17.]
[55]	Nan ZB ( 1996 c). Effects of Acremonium endophyte on the growth of Hordeum bogdanii. Pratacultural Science, 13, 16-18.
	[ 南志标 ( 1996 c). 内生真菌对布顿大麦草生长的影响. 草业科学, 13, 16-18.]
[56]	Nan ZB, Li CJ ( 2000). Neotyphodium in native grasses in China and observations on endophyte/host interaction. In: Paul VH, Dapprich PD eds. Proceedings of the 4th International Neotyphodium/Grass Interactions Symposium. Soest, Germany. 41-50.
[57]	Nan ZB, Li CJ ( 2004). Roles of the grass-Neotyphodium association in pastoral agriculture systems. Acta Ecologica Sinica, 24, 605-616. DOI URL
	[ 南志标, 李春杰 ( 2004). 禾草内生真菌在草地农业系统中的作用. 生态学报, 24, 605-616.] DOI URL
[58]	Nan ZB, Wang SM, Wang YR, Fu H, Li CJ, Duan TY ( 2016). Stress tolerance mechanisms of 6 native plant species growing in China’s northern grassland and their utilization. Chinese Science Bulletin, 61, 239-249.
	[ 南志标, 王锁民, 王彦荣, 傅华, 李春杰, 段廷玉 ( 2016). 我国北方草地6种乡土植物抗逆机理与应用. 科学通报, 61, 239-249.]
[59]	Norton BW, Li XS, Fletcher I ( 2009a). The grazing preferences of sheep on pastures containing drunken horse grass (Achnatherum inebrians) in Xinjiang Province, China. In: Papachristou TG, Parisi ZM, Salem HB, Morand-Fehr P eds. Twelfth Seminar of the Sub-Network FAO-CIHEAM on Sheep and Goat Nutrition. Thessaloniki, Greece. 105-110.
[60]	Norton BW, Li XS, Fletcher I ( 2009b). The effects of endophyte content on the nutritive value of drunken horse grass (Achnatherum inebrians) fed to sheep in Xinjiang Province, China. In: Papachristou TG, Parisi ZM, Salem HB, Morand-Fehr P eds. Twelfth Seminar of the Sub-Network FAO-CIHEAM on Sheep and Goat Nutrition. Thessaloniki, Greece. 147-152. DOI URL
[61]	Pennell CGL, Rolston MP, van Koten C, Hume DE, Card SD ( 2017). Reducing bird numbers at New Zealand airports a unique endophyte product. New Zealand Plant Protection, 70, 224-234. DOI URL
[62]	Qian SY, Zhang ZL ( 1986). Control of Achnatherum inebrians by dalapon in grassland. Gansu Animal and Veterinary Sciences, ( 4), 49.
	[ 钱顺云, 张正礼 ( 1986). 茅草枯灭除草原醉马草试验. 甘肃畜牧兽医, ( 4), 49. ]
[63]	Ren JZ ( 1954). Several common poisonous weeds in the northwestern grasslands. Animal Husbandry & Veterinary Medicine, ( 2), 56-60.
	[ 任继周 ( 1954). 西北草原上几种常见的毒草. 畜牧与兽医, ( 2), 56-60.]
[64]	Ren JZ ( 2013). China’s historical shifting between grassland-agriculture and cropland-agriculture. Agricultural History of China, 32(1), 3-8.
	[ 任继周 ( 2013). 中国草地农业系统与耕地农业系统的历史嬗替. 中国农史, 32(1), 3-8.]
[65]	Ren JZ, Jia ZZ ( 1961). Effect of burning on the Achnatherum inebrians. Journal of Gansu Agricultural University, ( 2), 1-5.
	[ 任继周, 贾宗周 ( 1961). 焚烧对醉马草丛的效果. 甘肃农业大学学报, ( 2), 1-5.]
[66]	Sahedula HM, Nuerlan, Jianate, Ma KR, Zhang JL, Guo XZ, Hali, Wu CY, Li SG, Wan GF, Zhen BL, Wang F, Wu DC, Zhu JY ( 1996). Drunken horse grass paper and manufacture method: China, ZL95118010.X. 1996-10-30.
	[ 萨赫都拉·霍曼, 努尔兰, 加娜特, 马克让, 张江玲, 郭选政, 哈里, 吴沧阳, 李世国, 万国富, 真标林, 王甫, 吴东成, 朱俊义 ( 1996). 醉马草纸及其制作方法: 中国, ZL95118010.X. 1996-10-30.]
[67]	Saimilakezi T, Jin GL, Zhang Y, Caidengba J, Chui GY, Hubisihaletu ( 2015). Study on allelopathic effect of Achnatherum inebrians on several forage. Xinjiang Agricultural Sciences, 52, 1715-1722. DOI URL
	[ 赛米拉克孜·台外库力, 靳瑰丽, 张勇娟, 才登巴·金保, 崔国盈, 胡毕斯哈勒图 ( 2015). 醉马草对几种牧草化感作用的研究. 新疆农业科学, 52, 1715-1722.] DOI URL
[68]	Shi DS, Ma RC, Han XZ, Bibulaai, Daerjia, Jiang QQ ( 1989). Report on control of Achnatherum inebrians. Journal of Xinjiang Agricultural University, ( 2), 7-12.
	[ 石定燧, 马瑞昌, 韩新忠, 比布拉艾, 达尔加, 蒋全勤 ( 1989). 禾本科醉马草防除试验报告. 新疆农业大学学报, ( 2), 7-12.]
[69]	Shi ZC ( 1997). Important Toxic Plants in Chinese Grasslands. China Agriculture Press, Beijing. 166-176.
	[ 史志诚 ( 1997). 中国草地重要有毒植物. 中国农业出版社, 北京. 166-176.]
[70]	Shymanovich T, Charlton ND, Musso AM, Scheerer J, Cech NB, Faeth SH, Young CA ( 2017). Interspecific and intraspecific hybrid Epichloë species symbiotic with the North American native grass Poa alsodes. Mycologia, 109, 459-474. DOI URL PMID
[71]	Siegel MR, Latch GCM, Johnson MC ( 1987). Fungal endophytes of grasses. Annal Revivew of Phytopathology, 25, 293-315. DOI URL
[72]	Sulitanghan ( 2010). Research on threaten and control of Achnatherum inebrians. Journal of Animal Industry in Xinjiang, ( 4), 56. DOI URL
	[ 苏里唐汗 ( 2010). 关于草地醉马草的危害及防除办法的探讨. 新疆畜牧业, ( 4), 56.] DOI URL
[73]	Sun YD, Zhang XX, Gu LJ, Li XZ, Wang P, Li CJ ( 2015). Antifungal activity of the crude extraction of endophyte-infected and endophyte-free drunken horse grass. Pratacultural Science, 32, 508-514. DOI URL
	[ 孙一丹, 张兴旭, 古丽君, 李秀璋, 王萍, 李春杰 ( 2015). 醉马草-内生真菌共生体中生物碱的抑菌活性. 草业科学, 32, 508-514.] DOI URL
[74]	Wan ZW ( 2017). Effects of Temperature, Illumination and pH Factor on Growth and Contents of Ergot Alkaloids of Epichloë Endophyte Symbiotic Achnatherum inebrians. Master degree dissertation, Lanzhou University, Lanzhou.
	[ 万志文 ( 2017). 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. 硕士学位论文, 兰州大学, 兰州.]
[75]	Wan ZW, Fen JR, Wang P, Li CJ ( 2016 a). Ergot alkaloids content of symbiont of Epichloë gansuensis-Achnatherum inebrians under different Mn, Zn and Fe condition. Acta Botanica Boreali-Occidentalia Sinica, 36, 1427-1434. DOI URL
	[ 万志文, 冯疆蓉, 王萍, 李春杰 ( 2016 a). Mn、Zn和Fe对醉马草内生真菌共生体麦角酰胺和麦角新碱含量的影响. 西北植物学报, 36, 1427-1434.] DOI URL
[76]	Wan ZW, Wang P, Zhang XX, Li CJ ( 2016 b). Effects of manganese and SA on germination of Achnatherum inebrians seed containing or free of endophyte. Acta Agrestia Sinica, 24, 107-113. DOI URL
	[ 万志文, 王萍, 张兴旭, 李春杰 ( 2016 b). 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, 24, 107-113.] DOI URL
[77]	Wang CS, Gao ZK, Xu WY ( 2007). The investigation of drunken horse grass in Xianqian town, Gaize County. Journal of Tibet Science and Technology, ( 3), 31-33.
	[ 王缠石, 高志楷, 徐文勇 ( 2007). 改则县先遣乡醉马草调研报告. 西藏科技, ( 3), 31-33.]
[78]	Wang JF, Nan ZB, Christensen MJ, Li CJ ( 2018 a). Glucose-6-phosphate dehydrogenase plays a vital role in Achnatherum inebrians plants host to Epichloë gansuensis by improving growth under nitrogen deficiency. Plant and Soil, 430, 37-48.
[79]	Wang JF, Nan ZB, Christensen MJ, Zhang XX, Tian P, Zhang ZX, Niu XL, Gao P, Chen TX, Ma LX ( 2018 b). Effect of Epichloë gansuensis endophyte on the nitrogen metabolism, nitrogen use efficiency, and stoichiometry of Achnatherum inebrians under nitrogen limitation. Journal of Agricultural and Food Chemistry, 66, 4022-4031.
[80]	Wang K, Dang XP ( 1991). Poisoning test of Achnatherum inebrians on sheep. Chinese Journal of Veterinary Science and Technology, 21(7), 32-33.
	[ 王凯, 党晓鹏 ( 1991). 醉马草对羊的毒性试验. 中国兽医科技, 21(7), 32-33.]
[81]	Wang L, Liao F, Huang GM, Liu YT, Luo JF, Zhou Q ( 2011). Double-colored real-time fluorescence PCR method for detection of Neotyphodium gansuense from drunken horse grass(Achnatherum inebrians). Journal of Agricultural Biotechnology, 19, 973-980.
	[ 汪琳, 廖芳, 黄国明, 刘跃庭, 罗加凤, 周琦 ( 2011). 双色荧光PCR检测醉马草内生真菌(Neotyphodium gansuense). 农业生物技术学报, 19, 973-980.]
[82]	Wang P ( 2015). Effects of Microelement and Exogenous Hormone on Growth and Ergot Alkaloid Content Achnatherum inebrians-Epichloë gansuensis Symbiont. Master degree dissertation, Lanzhou University, Lanzhou.
	[ 王萍 ( 2015). 微量元素和外源激素对醉马草内生真菌共生体生长及麦角生物碱含量的影响. 硕士学位论文, 兰州大学, 兰州.]
[83]	Wang P, Zhang XX, Zhao XJ, Li CJ ( 2014). Effects of ethphon and salicylic acid on growth and ergot alkaloids concentrations of Achnatherum inebrians seedlings. Pratacultural Science, 31, 2113-2118. DOI URL
	[ 王萍, 张兴旭, 赵晓静, 李春杰 ( 2014). 乙烯利及水杨酸对醉马草幼苗生长及生物碱含量的影响. 草业科学, 31, 2113-2118.] DOI URL
[84]	Welty RE, Milbrath GM, Faulkenberry D, Azevedo MD, Meek L, Hall K ( 1986). Endophyte detection in tall fescue seed by staining and ELISA. Seed Science and Technology, 14, 105-116. DOI URL
[85]	White JF, Morgan-Jones G, Morrow AC ( 1993). Taxonomy, life cycle, reproduction and detection of Acremonium endophytes. Agriculture, Ecosystems and Environment, 44, 13-37. DOI URL
[86]	Xia C, Christensen MJ, Zhang XX, Nan ZB ( 2018). Effect of Epichloë gansuensis endophyte and transgenerational effects on the water use efficiency, nutrient and biomass accumulation of Achnatherum inebrians under soil water deficit. Plant and Soil, 424, 555-571.
[87]	Xia C, Li N, Zhang XX, Feng Y, Christensen MJ, Nan ZB ( 2016). 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, 22, 26-34.
[88]	Xia C, Zhang XX, Christensen MJ, Nan ZB, Li CJ ( 2015). Epichloë endophyte affects the ability of powdery mildew(Blumeria graminis) to colonise drunken horse grass (Achnatherum inebrians). Fungal Ecology, 16, 26-33.
[89]	Xia C, Zhong R, Zhang XX, Nan ZB ( 2015). Allelopathic effects of volatile compounds from endophte-free and infected Achnatherum inebrians on Lolium perenne. Pratacultural Science, 32, 658-666. DOI URL
	[ 夏超, 钟睿, 张兴旭, 南志标 ( 2015). 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, 32, 658-666.] DOI URL
[90]	Xinjiang August 1st Agricultural College ( 1979) August 1st Agricultural College(1979). Identification Key of Plants in Xinjiang. Vol. 1. Xinjiang People’s Publishing House, ürümqi. 253.
	[ 新疆八一农学院 ( 1979). 新疆植物检索表(第一册). 新疆人民出版社, 乌鲁木齐. 253.]
[91]	Yan SL, Ba H, Haji AA ( 2004). Studies on the toxic constituent of Achnatherum inebrians. Natural Products Research and Development, 16, 395-398. DOI URL
	[ 颜世利, 巴杭, 阿吉艾克拜尔 ( 2004). 新疆醉马草化学成分的研究. 天然产物研究与开发, 16, 395-398.] DOI URL
[92]	Yang HL, Song YB, Sun ZJ, Jin GL, An SZ, Shi ZM, Ayiguli A ( 2015). Effects of different reseeding patterns on population characteristics of Achnatherum inebrians and diversity of grassland community. Guizhou Agricultural Sciences, 43(10), 67-71.
	[ 杨合龙, 宋跃斌, 孙宗玖, 靳瑰丽, 安沙舟, 石智明, 阿义古力·艾斯达吾列提 , ( 2015). 牧草不同补播方式对醉马草种群特征及其草地群落多样性的影响. 贵州农业科学, 43(10), 67-71.]
[93]	Yang S, Huang X, Chai Q, Li CJ, Nan ZB ( 2010). Allelopathic effects of endophytic fungi of Achnatherum inebrians Keng on the seed and seedling of three turf grasses. Acta Agrestia Sinica, 18, 78-83. DOI URL
	[ 杨松, 黄玺, 柴青, 李春杰, 南志标 ( 2010). 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, 18, 78-83.] DOI URL
[94]	Yao HC, Li XZ, Yang DQ, Duoji DZ ( 2007). The investigation on the hazard of drunken horse grass in Gaize County. Journal of Tibet Science and Technology, ( 2), 29-31. DOI URL
	[ 姚海潮, 李晓忠, 杨德全, 多吉顿珠 ( 2007). 改则县醉马草危害情况调研报告. 西藏科技, ( 2), 29-31.] DOI URL
[95]	Yao X ( 2015). The Ecological Regulating Effect of Achnatherum inebrians-Epichloë gansuensis Symbiont on Biodiversity and Diseases. Master degree dissertation, Lanzhou University, Lanzhou.
	[ 姚祥 ( 2015). 醉马草内生真菌共生体对草地生物多样性和病害的生态调控作用. 硕士学位论文, 兰州大学, 兰州.]
[96]	Yao X, Christensen MJ, Bao G, Zhang C, Li X, Li C, Nan Z ( 2015). A toxic endophyte-infected grass helps reverse degradation and loss of biodiversity of over-grazed grasslands in northwest China. Scientific Reports, 5, 18527. DOI: 10.1038/srep18527. DOI URL PMID
[97]	Yao X, Fan Y, Chai Q, Johnson RD, Nan ZB, Li CJ ( 2016). Modification of susceptible and toxic herbs on grassland disease. Scientific Reports, 7, 30635. DOI: 10.1038/srep30635. DOI URL
[98]	Zhang XX ( 2008). Effects of Endophyte Infection on Pest Resistance to Drunken Horse Grass. Master degree dissertation, Lanzhou University, Lanzhou.
	[ 张兴旭 ( 2008). 内生真菌对醉马草抗虫性的影响研究. 硕士学位论文. 兰州大学, 兰州.]
[99]	Zhang XX ( 2012). Response of Achnatherum inebrians/Neotyphodium gansuense Symbiont to Stress and Secondary Metabolities Activities. PhD dissertation, Lanzhou University, Lanzhou.
	[ 张兴旭 ( 2012). 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. 博士学位论文, 兰州大学, 兰州.]
[100]	Zhang XX, Fan XM, Li CJ, Nan ZB ( 2010 a). Effects of cadmium stress on seed germination, seedling growth and antioxidative enzymes in Achnatherum inebrians plants infected with a Neotyphodium endophyte. Plant Growth Regulation, 60, 91-97.
[101]	Zhang XX, Li CJ, Nan ZB ( 2010 b). Effects of cadmium stress on growth and anti-oxidative systems in Achnatherum inebrians symbiotic with Neoytphodium gansuense. Journal of Hazardous Materials, 175, 703-709.
[102]	Zhang XX, Li CJ, Nan ZB ( 2011). Effects of cutting frequency and height on alkaloid production in endophyte-infected drunken horse grass (Achnatherum inebrians). Science China: Life Science, 41, 422-426.
	[ 张兴旭, 李春杰, 南志标 ( 2011). 刈割高度和频度对醉马草(Achnatherum inebrians)内生真菌共生体麦角生物碱的影响. 中国科学: 生命科学, 41, 422-426.]
[103]	Zhang XX, Li CJ, Nan ZB ( 2011). Effects of salt and drought stress on alkaloid production in endophyte-infected drunken horse grass (Achnatherum inebrians). Biochemical Systematics and Ecology, 39, 471-476. DOI URL
[104]	Zhang XX, Li CJ, Nan ZB, Matthew C ( 2012). Neotyphodium endophyte increases Achnatherum inebrians(drunken horse grass) resistance to herbivores and seed predators. Weed Research, 52, 70-78.
[105]	Zhang XX, Nan ZB, Li CJ, Gao K ( 2014 a). Cytotoxic effect of ergot alkaloids in Achnatherum inebrians infected by the Neotyphodium gansuense endophyte. Journal of Agricultural & Food Chemistry, 62, 7419-7422.
[106]	Zhang XX, Wu YP, Nan ZB ( 2014 b). Antifungal activity of petroleum ether extracts from Achnatherum inebrians infected with Neotyphodium gansuense. Science China: Life Sciences, 57, 1234-1235.
[107]	Zhang YJ, Zhu ZQ ( 1984). Studies on the chemical compositions of Achnatherum inebrians. Chemical Journal of Chinese Universities, 3(S1), 150-152. DOI URL
	[ 张友杰, 朱子清 ( 1984). 醉马草化学成分的研究. 高等学校化学学报, 3(S1), 150-152.] DOI URL
[108]	Zhao XJ ( 2015). Studies on Detecting Methods and Microstructure of Epichloë endophytes of Grasses. Master degree dissertation, Lanzhou University, Lanzhou.
	[ 赵晓静 ( 2015). 禾草内生真菌显微结构及其检测方法的研究. 硕士学位论文, 兰州大学, 兰州.]
[109]	Zhong R, Xia C, Ju YW, Li NN, Zhang XX, Nan ZB, Christensen MJ ( 2018). Effects of Epichloë gansuensis on root-associated fungal communities of Achnatherum inebrians under different growth conditions. Fungal Ecology, 31, 29-36.

指标 Index	新疆 Xinjiang	青海 Qinghai	甘肃 Gansu	内蒙古 Nei Mongol	宁夏 Ningxia	西藏 Xizang
年降水量 Annual precipitation (mm)	124-277	282-450	129-558	213-300	286-323	262-977
海拔 Elevation (m)	1 200-2 200	2 600-3 910	1 647-3 056	1 750-2 090	1 883-1 937	3 028-4 305
纬度 Latitude (N)	41.78°-47.97°	34.15°-37.28°	34.77°-38.83°	38.90°-39.02°	36.48°-36.55°	29.48°-30.17°
经度 Longitude (E)	74.82°-94.82°	99.77°-101.77°	99.62°-105.20°	105.83°-105.90°	105.67°-105.72°	96.07°-97.28°
带菌率 Infection rate (%)	97.7 (83-100)	98.3 (75-100)	98.9 (80-100)	95.8 (80-100)	100	97.0 (0-100)

指标 Index	新疆 Xinjiang	青海 Qinghai	甘肃 Gansu	内蒙古 Nei Mongol	宁夏 Ningxia	西藏 Xizang
年降水量 Annual precipitation (mm)	124-277	282-450	129-558	213-300	286-323	262-977
海拔 Elevation (m)	1 200-2 200	2 600-3 910	1 647-3 056	1 750-2 090	1 883-1 937	3 028-4 305
纬度 Latitude (N)	41.78°-47.97°	34.15°-37.28°	34.77°-38.83°	38.90°-39.02°	36.48°-36.55°	29.48°-30.17°
经度 Longitude (E)	74.82°-94.82°	99.77°-101.77°	99.62°-105.20°	105.83°-105.90°	105.67°-105.72°	96.07°-97.28°
带菌率 Infection rate (%)	97.7 (83-100)	98.3 (75-100)	98.9 (80-100)	95.8 (80-100)	100	97.0 (0-100)