Advances in Alkaloid Biosynthesis and Regulation in Grass-Endophyte Symbioses

doi:10.11733/j.issn.1007-0435.2026.10.001

Acta Agrestia Sinica ›› 2026, Vol. 34 ›› Issue (10): 3525-3536.DOI: 10.11733/j.issn.1007-0435.2026.10.001

Advances in Alkaloid Biosynthesis and Regulation in Grass-Endophyte Symbioses

LIANG Zheng^1,2, WEI Xue-kai^1,2, YAN Wei-mo^1,2, WANG Jia-kun^1,2, LI Chun-jie^1,2,3

1. State Key Laboratory of Herbage Improvement and Grassland Agro-ecosystems/Key Laboratory of Grassland Livestock Industry Innovation, Ministry of Agriculture and Rural Affairs, Lanzhou University/Engineering Research Center of Grassland Agriculture and Ministry of Education, Lanzhou University/Western Grassland Technology Innovation Center of Gansu Province/Center for Grassland Microorganisms of Lanzhou University/College of Pastoral Agriculture Science and Technology of Lanzhou University, Lanzhou, Gansu Province 730020, China;
2. Jingtai Field Scientific Observation and Research Station of Pastoral Agriculture, Baiyin, Gansu Province 730900, China;
3. Grassland Research Center of National Forestry and Grassland Administration, Chinese Academy of Forestry, Beijing 100091, China

Received:2026-05-24 Revised:2026-07-14 Published:2026-10-10

禾草内生真菌共生体生物碱合成与调控的研究进展

梁政^1,2, 魏学凯^1,2, 闫伟茉^1,2, 王佳坤^1,2, 李春杰^1,2,3

1. 兰州大学草种创新与草地农业生态系统全国重点实验室/兰州大学农业农村部草牧业创新重点实验室/兰州大学草地农业教育部工程研究中心/甘肃省西部草业技术创新中心/兰州大学草地微生物研究中心/兰州大学草地农业科技学院, 甘肃兰州 730020;
2. 兰州大学景泰草地农业野外科学观测研究站, 甘肃白银 730900;
3. 中国林业科学研究院国家林业和草原局草原研究中心, 北京 100091

通讯作者: 李春杰, E-mail: chunjie@lzu.edu.cn
作者简介:梁政（2002-），男，汉族，河南安阳人，硕士研究生，主要从事牧草与草坪病理学方向研究，E-mail：liangzh2024@lzu.edu.cn
基金资助:
青年科学基金项目（C类）（32503291）；甘肃省青年科技基金（25JRRA698）资助

Abstract

Abstract: Alkaloids produced by Epichloë endophytes in grasses serve as an important chemical basis for mediating the symbiotic relationship’s insect-repellent effects and influencing the health of herbivores. Different alkaloids exhibit significant variations in their core structural scaffolds， biosynthetic pathways， tissue distribution， and biological activities. Their formation does not depend solely on the presence of biosynthetic genes， but is rather the result of the combined interactions among gene cluster composition， host genotype， and environmental conditions. This paper systematically compared the biosynthetic strategies and ecological effects of four major classes of alkaloids： peramine， ergot alkaloids， indole-diterpenes， and lolines. It further explored the multi-level regulatory mechanisms involving in alkaloid biosynthesis， including environmental factors， host signals， transcriptional regulation， and epigenetic regulation. Ultimately， this provides an important theoretical foundation for deeply understanding the functional mechanisms of the grass-endophyte symbiotic system and the agricultural applications of these alkaloids.

Key words: Grass endophyte, Epichloë, Alkaloid biosynthetic pathways, Secondary metabolism, Regulatory mechanism

摘要： 禾草Epichloë属内生真菌产生的生物碱是介导共生体抗虫效应及影响草食动物健康的重要化学基础。不同生物碱在核心骨架、合成途径、组织分布和生物活性方面具有显著差异，其形成并非单纯取决于合成基因的存在，而是基因簇组成、宿主基因型及环境条件共同作用的结果。本文系统比较了波胺、麦角碱、吲哚二萜和黑麦草碱四类主要生物碱的合成策略及其生态效应，进一步探讨了生物碱合成过程中的多层次调控机制，包括环境因子、宿主信号、转录调控与表观遗传调控等，为深入理解禾草内生真菌共生体系的功能机制及其生物碱的农业应用提供了重要的理论基础。

关键词: 禾草内生真菌共生体, Epichloë, 生物碱合成途径, 次生代谢, 调控机制

CLC Number:

S332.6

LIANG Zheng, WEI Xue-kai, YAN Wei-mo, WANG Jia-kun, LI Chun-jie. Advances in Alkaloid Biosynthesis and Regulation in Grass-Endophyte Symbioses[J]. Acta Agrestia Sinica, 2026, 34(10): 3525-3536.

梁政, 魏学凯, 闫伟茉, 王佳坤, 李春杰. 禾草内生真菌共生体生物碱合成与调控的研究进展[J]. 草地学报, 2026, 34(10): 3525-3536.

References

[1] NAN Z B， LI C J. Roles of the grass-Neotyphodium association in pastoral agriculture systems[J]. Ecological Research， 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[J]. 生态学报， 2004（3）： 605-616
[2] DU M X， WANG T， LI C J， et al. Advances in the taxonomy of the genus Epichloë endophytic fungi in grasses[J]. Acta Agrestia Sinica， 2023， 31（12）： 3575-3586 杜明祥，王添，李春杰，等. 禾草Epichloë属内生真菌分类学研究进展[J]. 草地学报， 2023， 31（12）： 3575-3586
[3] MWANGI N G， STEVENS M， WRIGHT A J D， et al. Grass-endophyte interactions and their associated alkaloids as a potential management strategy for plant parasitic nematodes[J]. Toxins， 2024， 16（6）： 274
[4] XIA C， CHRISTENSEN M J， ZHANG X， et al. Effect of Epichloë gansuensis endophyte and transgenerational effects on the water use efficiency， nutrient and biomass accumulation of Achnatherum inebrians under soil water deficit[J]. Plant and Soil， 2018， 424（1）： 555-571
[5] 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[J]. Plant Growth Regulation， 2016， 80（3）： 367-375
[6] XIA C， LI N N， ZHANG Y W， et al. Role of Epichloë endophytes in defense responses of cool-season grasses to pathogens： a review[J]. Plant Disease， 2018， 102（11）： 2061-2073
[7] SCHARDL C L， LEUCHTMANN A， SPIERING M J. Symbioses of grasses with seedborne fungal endophytes[J]. Annual Review of Plant Biology， 2004（55）： 315-340
[8] GARCÍA PARISI P A， GRIMOLDI A A， OMACINI M. Endophytic fungi of grasses protect other plants from aphid herbivory[J]. Fungal Ecology， 2014（9）： 61-64
[9] SCHARDL C L， YOUNG C A， HESSE U， et al. Plant-symbiotic fungi as chemical engineers： multi-genome analysis of the Clavicipitaceae reveals dynamics of alkaloid loci[J]. Plos Genetics， 2013， 9（2）： e1003323
[10] ZHANG W， FORESTER N T， MOON C D， et al. Epichloë seed transmission efficiency is influenced by plant defense response mechanisms[J]. Frontiers in Plant Science， 2022（13）： 1025698
[11] SHI Q. Effects of elevation on biological characteristics of Elymus tangutorum-Epichloë endophytic symbiont[D]. Lanzhou： Lanzhou University， 2021： 14-20 师茜. 海拔高度与麦宾草-内生真菌共生体生物学特性[D]. 兰州：兰州大学， 2021： 14-20
[12] YOUNG C A， HUME D E， MCCULLEY R L. Forages and pastures symposium： fungal endophytes of tall fescue and perennial ryegrass： pasture friend or foe [J]. Journal of Animal Science， 2013， 91（5）： 2379-2394
[13] TAYLOR A A， FIKE J H， MERCADANTE V R G， et al. The effects of endophyte-infected tall fescue on the production and reproductive performance parameters of beef cattle and calves[J]. Grasses， 2025， 4（3）： 29
[14] BACON C W， PORTER J K， ROBBINS J D， et al. Epichloë typhina from toxic tall fescue grasses[J]. Applied and Environmental Microbiology， 1977， 34（5）： 576-581
[15] BUSH L P， WILKINSON H H， SCHARDL C L. Bioprotective alkaloids of grass-fungal endophyte symbioses[J]. Plant Physiology， 1997， 114（1）： 1-7
[16] SONG Q Y， LI F， NAN Z B， et al. Do Epichloë endophytes and their grass symbiosis only produce toxic alkaloids to insects and livestock [J]. Journal of Agricultural and Food Chemistry， 2020， 68（5）： 1169-1185
[17] KOULMAN A， LANE G A， CHRISTENSEN M J， et al. Peramine and other fungal alkaloids are exuded in the guttation fluid of endophyte-infected grasses[J]. Phytochemistry， 2007， 68（3）： 355-360
[18] FUCHS B， KRISCHKE M， MUELLER M J， et al. Peramine and lolitrem B from endophyte-grass associations cascade up the food chain[J]. Journal of Chemical Ecology， 2013， 39（11）： 1385-1389
[19] KUANG Y. Characteristics of Epichloë endophyte-Festuca sinensis symbiote[D]. Lanzhou： Lanzhou University， 2016： 12-14 旷宇. 中华羊茅—内生真菌共生体的筛选与鉴定[D]. 兰州：兰州大学， 2016： 12-14
[20] SONG H. Phylogeny of nine Elymus species and related asexual Epichloë endophyte[D]. Lanzhou： Lanzhou University， 2016： 21-22 宋辉. 九种披碱草属植物及其Epichloё内生真菌的系统进化[D]. 兰州：兰州大学， 2016： 21-22
[21] KAUPPINEN M， SAIKKONEN K， HELANDER M， et al. Epichloë grass endophytes in sustainable agriculture[J]. Nature Plants， 2016， 2（2）： 15224
[22] RAHNAMA M， MACLEAN P， FLEETWOOD D J， et al. VelA and LaeA are key regulators of Epichloë festucae transcriptomic response during symbiosis with perennial ryegrass[J]. Microorganisms， 2020， 8（1）： 33
[23] XU W B. Study on the taxonomy， diversity andmetabolites of Epichloë endophyte in Festuca sinensis[D]. Lanzhou： Lanzhou University， 2023： 16-20 许文博. 中华羊茅内生真菌的分类、多样性及代谢产物研究[D]. 兰州：兰州大学， 2023： 16-20
[24] DU M X， WANG T， LI C J， et al. Discovery and characterization of Epichloë fungal endophytes from Elymus spp. in northwest China[J]. Microorganisms， 2024， 12（7）： 1497
[25] JIN Y Y. Microbial mechanisms of nitrogen effective use in Achnatherum inebrians-fungal endphyte symbiont[D]. Lanzhou： Lanzhou University， 2025： 15-24 金媛媛. 醉马草-内生真菌共生体氮高效利用的微生物机制[D]. 兰州：兰州大学， 2025： 15-24
[26] MOORE J R， PRATLEY J E， MACE W J， et al. Variation in alkaloid production from genetically diverse lolium accessions infected with Epichloë species[J]. Journal of Agricultural and Food Chemistry， 2015， 63（48）： 10355-10365
[27] SHIBA T， SUGAWARA K. Fungal loline alkaloids in grass–endophyte associations confer resistance to the rice leaf bug， Trigonotylus caelestialium[J]. Entomologia Experimentalis et Applicata， 2009， 130（1）： 55-62
[28] GE H M， TAN R X. Symbionts， an important source of new bioactive natural products[J]. Progress in Chemistry， 2009， 21（1）： 30-46 戈惠明，谭仁祥. 共生菌——新活性天然产物的重要来源[J]. 化学进展， 2009， 21（1）： 30-46
[29] PANACCIONE D G， BEAULIEU W T， COOK D. Bioactive alkaloids in vertically transmitted fungal endophytes[J]. Functional Ecology， 2014， 28（2）： 299-314
[30] VASSILIADIS S， REDDY P， HEMSWORTH J， et al. Quantitation and distribution of Epichloë-derived alkaloids in perennial ryegrass tissues[J]. Metabolites， 2023， 13（2）： 205
[31] ROWAN D D， HUNT M B， GAYNOR D L. Peramine， a novel insect feeding deterrent from ryegrass infected with the endophyte Acremonium loliae[J]. Journal of the Chemical Society， Chemical Communications， 1986， 22（12）： 935-936
[32] TANAKA A， TAPPER B A， POPAY A， et al. A symbiosis expressed non-ribosomal peptide synthetase from a mutualistic fungal endophyte of perennial ryegrass confers protection to the symbiotum from insect herbivory[J]. Molecular Microbiology， 2005， 57（4）： 1036-1050
[33] LUO H P， XIE L X， ZENG J， et al. Biosynthesis and regulation of bioprotective alkaloids in the gramineae endophytic fungi with implications for herbivores deterrents[J]. Current Microbiology， 2015， 71（6）： 719-724
[34] WANG Y Q， LIU H J， LI H， et al. Research progress on the production and biosynthesis of ergotamine[J]. West China Journal of Pharmaceutical Sciences， 2015， 30（1）： 109-112 王雅琴，刘红娟，李海，等. 麦角碱的生产及生物合成的研究进展[J]. 华西药学杂志， 2015， 30（1）： 109-112
[35] WALLWEY C， LI S M. Ergot alkaloids： structure diversity， biosynthetic gene clusters and functional proof of biosynthetic genes[J]. Natural Product Reports， 2011， 28（3）： 496-510
[36] PANACCIONE D G， RYAN K L， SCHARDL C L， et al. Chapter twelve - Analysis and modification of ergot alkaloid profiles in fungi[M] Pittsburgh： Academic Press， 2012： 267-290
[37] WANG T. Screening of novel Epichloë endophyte strains and their metabolites analyses[D]. Lanzhou： Lanzhou University， 2023： 6-10 王添. 禾草Epichloë属内生真菌优良菌株筛选及代谢产物分析[D]. 兰州：兰州大学， 2023： 6-10
[38] EASTON H S， LATCH G C M， TAPPER B A， et al. Ryegrass host genetic control of concentrations of endophyte-derived alkaloids[J]. Crop Science， 2002， 42（1）： 51-57
[39] PHILIPPE G. Lolitrem B and indole diterpene alkaloids produced by endophytic fungi of the genus Epichloë and their toxic effects in livestock[J]. Toxins， 2016， 8（2）： 47
[40] PETROSKI R， YATES S， WEISLEDER D， et al. Isolation， semi-synthesis， and Nmr spectral studies of loline alkaloids[J]. Journal of Natural Products， 2004， 52（4）： 810-817
[41] FINKING R， MARAHIEL M A. Biosynthesis of nonribosomal peptides[J]. Annual Review of Microbiology， 2004（58）： 453-488
[42] KELLER N P， TURNER G， BENNETT J W. Fungal secondary metabolism-from biochemistry to genomics[J]. Nature Reviews Microbiology， 2005， 3（12）： 937-947
[43] HAARMANN T， ORTEL I， TUDZYNSKI P， et al. Identification of the cytochrome P450 monooxygenase that bridges the clavine and ergoline alkaloid pathways[J]. ChemBioChem， 2006， 7（4）： 645-652
[44] LORENZ N， WILSON E V， MACHADO C， et al. Comparison of ergot alkaloid biosynthesis gene clusters in Claviceps species indicates loss of late pathway steps in evolution of C. fusiformis[J]. Applied and Environmental Microbiology， 2007， 73（22）： 7185-7191
[45] GEBLER J C， POULTER C D. Purification and characterization of dimethylallyl tryptophan synthase from Claviceps purpurea[J]. Archives of Biochemistry and Biophysics， 1992， 296（1）： 308-313
[46] SCHARDL C L， PANACCIONE D G， TUDZYNSKI P. Chapter 2 Ergot alkaloids - biology and molecular biology[J]. The Alkaloids. Chemistry and Biology， 2006（63）： 45-86
[47] GERHARDS N， NEUBAUER L， TUDZYNSKI P， et al. Biosynthetic pathways of ergot alkaloids[J]. Toxins， 2014， 6（12）： 3281-3295
[48] NIELSEN C A， FOLLY C， HATSCH A， et al. The important ergot alkaloid intermediate chanoclavine-I produced in the yeast Saccharomyces cerevisiae by the combined action of EasC and EasE from Aspergillus japonicus[J]. Microbial Cell Factories， 2014， 13（1）： 95
[49] RIGBERS O， LI S M. Ergot alkaloid biosynthesis in Aspergillus fumigatus[J]. Journal of Biological Chemistry， 2008， 283（40）： 26859-26868
[50] LORENZ N， OLŠOVSKÁ J， ŠULC M， et al. Alkaloid cluster gene ccsA of the ergot fungus Claviceps purpurea encodes chanoclavine I synthase， a flavin adenine dinucleotide-containing oxidoreductase mediating the transformation of N-methyl-dimethylallyltryptophan to chanoclavine I[J]. Applied and Environmental Microbiology， 2010， 76（6）： 1822-1830
[51] WALLWEY C， MATUSCHEK M， LI S M. Ergot alkaloid biosynthesis in Aspergillus fumigatus： conversion of chanoclavine-I to chanoclavine-I aldehyde catalyzed by a short-chain alcohol dehydrogenase FgaDH[J]. Archives of Microbiology， 2010， 192（2）： 127-134
[52] CHENG J Z， COYLE C M， PANACCIONE D G， et al. A role for Old Yellow Enzyme in ergot alkaloid biosynthesis[J]. Journal of the American Chemical Society， 2010， 132（6）： 1776-1777
[53] COYLE C M， CHENG J Z， O’CONNOR S E， et al. An Old Yellow Enzyme gene controls the branch point between Aspergillus fumigatus and Claviceps purpurea ergot alkaloid pathways[J]. Applied and Environmental Microbiology， 2010， 76（12）： 3898-3903
[54] CORREIA T， GRAMMEL N， ORTEL I， et al. Molecular cloning and analysis of the ergopeptine assembly system in the ergot fungus Claviceps purpurea[J]. Chemistry & Biology， 2003， 10（12）： 1281-1292
[55] HAARMANN T， LORENZ N， TUDZYNSKI P. Use of a nonhomologous end joining deficient strain （Δku70） of the ergot fungus Claviceps purpurea for identification of a nonribosomal peptide synthetase gene involved in ergotamine biosynthesis[J]. Fungal Genetics and Biology， 2008， 45（1）： 35-44
[56] NIU J W， QI J Z， WANG P C， et al. The chemical structures and biological activities of indole diterpenoids[J]. Natural Products and Bioprospecting， 2023， 13（1）： 3
[57] SCHARDL C L， YOUNG C A， FAULKNER J R， et al. Chemotypic diversity of Epichloë， fungal symbionts of grasses[J]. Fungal Ecology， 2012， 5（3）： 331-344
[58] TAGAMI K， LIU C W， MINAMI A， et al. Reconstitution of biosynthetic machinery for indole-diterpene paxilline in Aspergillus oryzae[J]. Journal of the American Chemical Society， 2013， 135（4）： 1260-1263
[59] YOUNG C A， BRYANT M K， CHRISTENSEN M J， et al. Molecular cloning and genetic analysis of a symbiosis-expressed gene cluster for lolitrem biosynthesis from a mutualistic endophyte of perennial ryegrass[J]. Molecular genetics and genomics： MGG， 2005， 274（1）： 13-29
[60] LUDLOW E J， VASSILIADIS S， EKANAYAKE P N， et al. Analysis of the indole diterpene gene cluster for biosynthesis of the epoxy-janthitrems in Epichloë endophytes[J]. Microorganisms， 2019， 7（11）： 560
[61] SAIKIA S， TAKEMOTO D， TAPPER B A， et al. Functional analysis of an indole-diterpene gene cluster for lolitrem B biosynthesis in the grass endosymbiont Epichloë festucae[J]. FEBS Letters， 2012， 586（16）： 2563-2569
[62] YOUNG C A， FELITTI S， SHIELDS K， et al. A complex gene cluster for indole-diterpene biosynthesis in the grass endophyte Neotyphodium lolii[J]. Fungal Genetics and Biology， 2006， 43（10）： 679-693
[63] BHARADWAJ R， JAGADEESAN H， KUMAR S R， et al. Molecular mechanisms in grass-Epichloë interactions： towards endophyte driven farming to improve plant fitness and immunity[J]. World Journal of Microbiology and Biotechnology， 2020， 36（7）： 92
[64] XU W， GAVIA D J， TANG Y. Biosynthesis of fungal indole alkaloids[J]. Natural Product Reports， 2014， 31（10）： 1474-1487
[65] MUNDAY-FINCH S C， WILKINS A L， MILES C O， et al. Isolation and structure elucidation of lolilline， a possible biosynthetic precursor of the lolitrem family of tremorgenic mycotoxins[J]. Journal of Agricultural and Food Chemistry， 1997， 45（1）： 199-204
[66] WILKINSON H H， SIEGEL M R， BLANKENSHIP J D， et al. Contribution of fungal loline alkaloids to protection from aphids in a grass-endophyte mutualism[J]. Molecular Plant-Microbe Interactions， 2007， 13（10）： 1027-1033
[67] SPIERING M J， MOON C D， WILKINSON H H， et al. Gene clusters for insecticidal loline alkaloids in the grass-endophytic fungus Neotyphodium uncinatum[J]. Genetics， 2005， 169（3）： 1403-1414
[68] KUTIL B L， GREENWALD C， LIU G， et al. Comparison of loline alkaloid gene clusters across fungal endophytes： predicting the co-regulatory sequence motifs and the evolutionary history[J]. Fungal Genetics and Biology， 2007， 44（10）： 1002-1010
[69] BLANKENSHIP J D， HOUSEKNECHT J B， PAL S， et al. Biosynthetic precursors of fungal pyrrolizidines， the loline alkaloids[J]. ChemBioChem， 2005， 6（6）： 1016-1022
[70] SPIERING M J， FAULKNER J R， ZHANG D X， et al. Role of the LolP cytochrome P450 monooxygenase in loline alkaloid biosynthesis[J]. Fungal Genetics and Biology， 2008， 45（9）： 1307-1314
[71] FAULKNER J R， HUSSAINI S R， BLANKENSHIP J D， et al. On the sequence of bond formation in loline alkaloid biosynthesis[J]. ChemBioChem， 2006， 7（7）： 1078-1088
[72] XU Y Q， LIU S N， GAO J M， et al. Molecular basis for Cγ-N bond formation by PLP-dependent enzyme LolC[J]. Biochemistry， 2024， 63（24）： 3348-3356
[73] PAN J， BHARDWAJ M， FAULKNER J R， et al. Ether bridge formation in loline alkaloid biosynthesis[J]. Phytochemistry， 2014， 98： 60-68
[74] PAN J， BHARDWAJ M， ZHANG B， et al. Installation of the ether bridge of lolines by the iron- and 2-oxoglutarate-dependent oxygenase， LolO： regio- and stereochemistry of sequential hydroxylation and oxacyclization reactions[J]. Biochemistry， 2018， 57（14）： 2074-2083
[75] PAN J， BHARDWAJ M， NAGABHYRU P， et al. Enzymes from fungal and plant origin required for chemical diversification of insecticidal loline alkaloids in grass-Epichloë symbiota[J]. PloS One， 2014， 9（12）： e115590
[76] SPIERING M J， WILKINSON H H， BLANKENSHIP J D， et al. Expressed sequence tags and genes associated with loline alkaloid expression by the fungal endophyte Neotyphodium uncinatum[J]. Fungal Genetics and Biology， 2002， 36（3）： 242-254
[77] CHUJO T， LUKITO Y， EATON C J， et al. Complex epigenetic regulation of alkaloid biosynthesis and host interaction by heterochromatin protein I in a fungal endophyte-plant symbiosis[J]. Fungal genetics and biology， 2019（125）： 71-83
[78] RUIZ-VILLAFÁN B， CRUZ-BAUTISTA R， MANZO-RUIZ M， et al. Carbon catabolite regulation of secondary metabolite formation， an old but not well-established regulatory system[J]. Microbial Biotechnology， 2022， 15（4）： 1058-1072
[79] BYLIN A， HUME D， CARD S， et al. Influence of nitrogen fertilization on growth and loline alkaloid production of meadow fescue （Festuca pratensis） associated with the fungal symbiont Neotyphodium uncinatum[J]. Botany， 2014， 92： 370-376
[80] JIANG W L， HOU X Y， PENG G X， et al. Fungal ergot alkaloids： metabolic pathways， biological functions， and advances in synthetic reprogramming[J]. Biotechnology Advances， 2025（81）： 108578
[81] MULINTI P， ALLEN N A， COYLE C M， et al. Accumulation of ergot alkaloids during conidiophore development in Aspergillus fumigatus[J]. Current Microbiology， 2014， 68（1）： 1-5
[82] RYAN K L， MOORE C T， PANACCIONE D G. Partial reconstruction of the ergot alkaloid pathway by heterologous gene expression in Aspergillus nidulans[J]. Toxins， 2013， 5（2）： 445-455
[83] UPPALA S S， WU B M， ALDERMAN S C. Effects of temperature and duration of preconditioning cold treatment on sclerotial germination of Claviceps purpurea[J]. Plant Disease， 2016， 100（10）： 2080-2086
[84] TIAN F， LEE S Y， WOO S Y， et al. Transcriptomic responses of Aspergillus flavus to temperature and oxidative stresses during aflatoxin production[J]. Scientific Reports， 2021， 11（1）： 2803
[85] LIN W H， GAO C F， WANG J J， et al. Effects of drought stress on peramine and lolitrem B in Epichloë-endophyte-infected perennial ryegrass[J]. Life， 2022， 12（8）： 1207
[86] SHI C， WANG C Z， HE J K， et al. Epichloë endophytes potentially facilitate host plant recruitment of rhizosphere microbiota carrying beneficial traits[J]. Physiologia Plantarum， 2025， 177（4）： e70397
[87] XU H E， HU M F， ZHANG B Y， et al. The effect of symbiosis between endophytic fungi and Hordeum Bogdanii on nutrients and enzyme activities in rhizosphere and non-rhizosphere Soils[J]. Acta Agrestia Sinica， 2025， 34（1）： 23-32 徐洪娥，户萌菲，张博阳，等. 内生真菌与布顿大麦共生对根际与非根际土壤养分和酶活性的影响[J]. 草地学报， 2025， 34（1）： 23-32
[88] LIU J， CHEN Z J， WHITE J F， et al. Ergot alkaloid and endogenous hormones quantities and relationship in Epichloë endophyte： drunken horse grass are affected by altitude[J]. Journal of Plant Growth Regulation， 2023， 42（3）： 1979-1990
[89] POPAY A J， MACE W J， FINCH S C， et al. Epichloë fungal endophyte strains and their lolium hosts affect resistance to Listronotus bonariensis （Coleoptera： Curculionidae）[J]. New Zealand Journal of Agricultural Research， 2025， 68（5）： 997-1016
[90] REALINI F M， ESCOBEDO V M， UENO A C， et al. Anti-herbivory defences delivered by Epichloë fungal endophytes： a quantitative review of alkaloid concentration variation among hosts and plant parts[J]. Annals of Botany， 2024， 133（4）： 509-520
[91] ZHANG G， YAN P， LENG D D， et al. Functional roles of LaeA-like genes in fungal growth， cellulase activity， and secondary metabolism in Pleurotus ostreatus[J]. Journal of Fungi， 2022， 8（9）： 902
[92] BOK J W， KELLER N P. LaeA， a regulator of secondary metabolism in Aspergillus spp[J]. Eukaryotic Cell， 2004， 3（2）： 527-535
[93] CHUJO T， SCOTT B. Histone H3K9 and H3K27 methylation regulates fungal alkaloid biosynthesis in a fungal endophyte-plant symbiosis[J]. Molecular Microbiology， 2014， 92（2）： 413-434
[94] LI D， LU P， YUAN B， et al. Research progress of endopfyte fungi from locoweed producing SW[J]. Acta Agrestia Sinica， 2024， 32（2）： 358-368 李丹，卢萍，袁博，等. 产苦马豆素的疯草内生真菌研究进展[J]. 草地学报， 2024， 32（2）： 358-368
[95] PENNELL C G L， POPAY A J， ROLSTON M P， et al. Avanex unique endophyte technology： reduced insect food source at airports[J]. Environmental Entomology， 2016， 45（1）： 101-108
[96] LIAN H N， CHEN T X， CHEN Z J， et al. Potential analysis of Achnatherum inebrians Epichloё endophyte infestation as an airfield grass to repel birds[J]. Acta Agrestia Sinica， 2021， 29（8）： 1649-1657 连鹤娜，陈泰祥，陈振江，等. 内生真菌醉马草作为机场驱鸟草的潜力分析[J]. 草地学报， 2021， 29（8）： 1649-1657
[97] GUO Y Z， ZHAO S J， SHA R K， et al. Study progress of poisonous plants containing pyrrolizidine alkaloids in natural grasslands[J]. Acta Agrestia Sinica， 2022， 30（2）： 297-311 郭亚洲，赵世姣，沙日扣，等. 天然草地含吡咯里西啶类生物碱有毒植物研究进展[J]. 草地学报， 2022， 30（2）： 297-311

Advances in Alkaloid Biosynthesis and Regulation in Grass-Endophyte Symbioses

禾草内生真菌共生体生物碱合成与调控的研究进展

PDF

Knowledge

Abstract

Cite this article

share this article

References

Related Articles 5

Recommended Articles

Metrics

Comments

[1]	CHEN Cai-jin, MA Lin, BAO Ming-fang, JIANG Qing-xue, ZHANG Guo-hui, ZHANG Shang-pei, GAO Ting, LIU Wen-hui, WANG Xue-min. Research Progress on the Role of WRKY Family Genes in Plants [J]. Acta Agrestia Sinica, 2025, 33(7): 2059-2069.
[2]	DU Ming-xiang, WANG Tian, LI Chun-jie, CHEN Tai-xiang. Advances in the Taxonomy of the Genus Epichloë Endophytic Fungi in Grasses [J]. Acta Agrestia Sinica, 2023, 31(12): 3575-3586.
[3]	LI Wen-qian, ZHANG Ying, WANG Meng-ru, CAO Jian, CHEN Jia-wei, CHAI Zhi, HOU Mei-li, LIANG Jian-ping. Effects of Different Fertilization Ratio on Yield and Quality of Astragalus membranaceus var.mongholicus [J]. Acta Agrestia Sinica, 2020, 28(1): 221-229.
[4]	GUO Yan-e, Li Ying-de, GAO Ping, WANG Zhi-gang, DUAN Ting-yu. Effects of Claroideoglomus etunicatum and Grass Endophyte on the Growth of Lolium perenne under Different Phosphorus Levels [J]. Acta Agrestia Sinica, 2018, 26(6): 1458-1466.
[5]	SONG Mei-ling, LI Chun-jie, PENG Qing-qing, LIANG Ying, NAN Zhi-biao. Effects of Neotyphodium Endophyte on Germination of Hordeum brevisubulatum under Temperature and Water Stress Conditions [J]. , 2010, 18(6): 833-837.