冬小包脚菇可培养内生真菌及菌基土壤真菌物种多样性分析

李鹏; 熊雪; 向准; 和耀威; 刘忠玄; 王晶

doi:10.19303/j.issn.1008-0384.2022.008.014

冬小包脚菇可培养内生真菌及菌基土壤真菌物种多样性分析

doi: 10.19303/j.issn.1008-0384.2022.008.014

贵州省生物研究所，贵州　贵阳　550009

基金项目: 贵州省科技计划项目（黔科合支撑[2019]2332号、黔科合支撑[2021]一般196、黔科合服企[2018]4002）

详细信息

作者简介:
李鹏（1987−），男，硕士，助理研究员，主要从事食用菌品种选育与栽培研究（E-mail：761410952@qq.com）

通讯作者:
向准（1976−），男，硕士，副研究员，主要从事食用菌品种选育与栽培研究（E-mail：xiangzhun@163.com）

中图分类号: TS 201.3
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出版历程
- 收稿日期: 2022-02-04
- 修回日期: 2022-05-19
- 网络出版日期: 2022-08-29
- 刊出日期: 2022-08-28

Diversity of Culturable Endophytes and Rhizosphere Fungi of Volvariella brumalis

Guizhou Institute of Biology, Guiyang, Guizhou　550027, China

摘要

摘要: 目的明确冬小包脚菇菌基土壤真菌和可培养内生真菌组成情况，为冬小包脚菇子实体分离及栽培奠定基础。方法以野生冬小包脚菇子实体及其菌基土壤为研究对象，采用传统组织分离法对子实体进行分离、纯化及ITS测序鉴定，并结合高通量测序法对菌基土壤真菌进行测序。结果分离得到的冬小包脚菇纯菌株属于Volvopluteus类群，也可属于传统定义的Volvariella类群。共得到冬小包脚菇内生真菌菌株50个，分属为3门13科16属20种，另有未定义科2个菌株，分别属于2个属，其中镰刀菌属（Fusarium）和毛霉属（Mucor）为优势属，且冬小包脚菇子实体内生真菌Shannon-Wiener多样性指数（H）为2.45，Simpson指数（Ｄ）为0.88，均匀度指数Pielou（J）为0.88，Margalef丰富度指数（R）为3.83。而冬小包脚菇菌基土壤中共检测到平均样品序列数67 919个，鉴定出276种土壤真菌，归属于10门131科213属，优势属为被孢霉属（Mortierella）、瓶毛壳属（Lophotrichus）、假散囊菌属（Pseudeurotium）、Pseudaleuria和两个未定义属unclassified_o_Sordariales、unclassified_c_Sordariomycetes。3个土样平均Ace指数达298.7，Shannon指数达3.53，Simpson指数为0.08，Chao 指数为304.23。结论冬小包脚菇可培养内生真菌和菌基土壤真菌都具有较为丰富的物种组成，该结论对冬小包脚菇菌种分离、培养、病虫害防治等栽培要点提供一定理论指导。
- 冬小包脚菇 /
- 鉴定 /
- 内生真菌 /
- 菌基土壤 /
- 物种组成
Abstract: Objective To promote the cold climate, specialty mushrooms in Guizhou, Volvariella brumalis as an ideal fill-in of the popular straw mushroom, V. volvacea, for wintertime market, the culturable endophytes and rhizosphere fungi associated with it were isolated and investigated to aid the cultivation. Methods Specimens from the fruiting bodies and rhizosphere soil of V. brumalis in the wild were collected. Endophytes in the fruiting bodies were isolated, cultured, and identified by ITS sequencing using the traditional tissue separation method, and fungal community in the rhizosphere soil analyzed by high-throughput sequencing. Results The 50 isolated endophytic strains were Volvopluteus, or the traditionally defined Volvariella, genera. They belonged to 3 phyla, 13 families, 16 genera (two of undefined families), and 20 species. Among them, Fusarium and Mucor were the dominant genera. The fungal Shannon-Wiener diversity index was 2.45, the Simpson index 0.88, the evenness index Pielou 0.88, and the Margalef richness index 3.83. In the rhizosphere soil, 67 919 sequences were detected with 276 kinds of fungi that belonged to 10 phyla, 131 families, and 213 genera. The dominant genera included Mortierella, Lophotricus, Pseudeurotium, Pseudaleuria, unclassified_o_Sordariales, and unclassified_c_Sordariomycetes. The Ace indices of 3 soil samples averaged 298.7, the Shannon indices 3.53, the Simpson indices 0.08, and the Chao indices 304.23. Conclusion Both the culturable endophytes and the rhizosphere fungi of V. brumalis were relatively rich in diversity. The information obtained would facilitate further research on the isolation, cultivation, and pest control of the mushroom for potentially valuable marketing in winter.
- Volvariella brumalis /
- identification /
- endophytic fungi /
- rhizosphere soil /
- fungal diversity

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图 1 基于ITS序列构建的冬小包脚菇系统发育树

Figure 1. Phylogenetic tree of V. brumalis constructed based on ITS sequence

下载: 全尺寸图片幻灯片

图 2 基于属水平的冬小包脚菇菌基土壤真菌物种组成

Figure 2. Composition of V. brumalis rhizosphere soil fungi at genus level

下载: 全尺寸图片幻灯片

表 1 冬小包脚菇可培养内生真菌菌群组成

Table 1. Composition of culturable endophytes in V. brumalis

门 Phylum	科 Family	属 Genus	菌株数 Number of strains	相对丰度 Relative abundance/%	相似种 Similar species	菌株数 Number of strains
担子菌门 Basidomycota	原毛平革菌科 Phanerochaetaceae	烟管菌属 Bjerkandera	3	6.00	烟管菌 Bjerkandera adusta	3
	球盖菇科 Strophariaceae	球盖菇属 Stropharia	4	8.00	大球盖菇 Stropharia rugosoannulata	4
	皱孔菌科 Meruliaceae	射脉菌属 Phlebia	1	2.00	胶质射脉革菌 Phlebia tremellosa	1
	多孔菌科 Polyporales	多孔菌属 Polyporus	2	4.00	漏斗多孔菌 Polyporus arcularius	2
	裂褶菌科 Schizophyllaceae	裂褶菌属 Schizophyllum	3	6.00	裂褶菌 Schizophyllum commune	3
接合菌门 Zygomycota	毛霉菌科 Mucoraceae	毛霉属 Mucor	8	16.00	不规则毛霉 Mucor irregularis	2
接合菌门 Zygomycota	毛霉菌科 Mucoraceae	毛霉属 Mucor	8	16.00	冻土毛霉 Mucor hiemalis	6
子囊菌门 Ascomycota	丛赤壳科 Nectriaceae	镰刀菌属 Fusarium	12	24.00	禾谷镰刀菌 Fusarium graminearum	6
					木贼镰刀菌 Fusarium equiseti	3
					尖孢镰刀菌 Fusarium oxysporum	3
	瘤座孢科 Tubeculariaceae	漆斑菌属 Myrothecium	1	2.00	Myrothecium sp.	1
	生赤壳科 Bionectriaceae	粘帚霉菌属 Clonostachys	3	6.00	粉红粘帚菌 Clonostachys rosea	3
	梨孢假壳科 Apiosporaceae	节菱孢属 Arthrinium	3	6.00	暗孢节菱孢菌 Arthrinium phaeospermum	3
	革菌科 Coriolaceae	蜡孔菌属 Ceriporia	1	2.00	撕裂蜡孔菌 Ceriporia lacerata	1
	曲霉科 Aspergillaceae	青霉属 Penicillium	1	2.00	Penicillium aquaticum	1
		曲霉属 Aspergillus	3	6.00	烟曲霉 Aspergillus fumigatus	1
		曲霉属 Aspergillus	3	6.00	聚多曲霉 Aspergillus sydowii	2
	山野壳菌科 Montagnulaceae	小球壳孢属 Microsphaeropsis	3	6.00	Microsphaeropsis amaranthi	3
	未定义	Leiothecium	1	2.00	Leiothecium ellipsoideum	1
	未定义	Parengyodontium	1	2.00	Parengyodontium album	1

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表 2 Alpha多样性指数分析

Table 2. Alpha diversity index analysis

土壤样品 Soil sample	序列数 Number	Ace指数 Ace index	Shannon指数 Shannon index	Simpson指数 Simpson index	Chao指数 Chao index	覆盖度 Coverage/%
S1	60290.00	334.90	3.11	0.12	341.14	99.88
S2	71619.00	283.08	4.05	0.04	292.00	99.95
S3	71849.00	278.25	3.44	0.08	279.56	99.94
平均值 Mean	67919.33	298.74	3.53	0.08	304.23	—

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参考文献(49)

[1]	何绍昌, 冯丽. 小包脚菇属一新种 [J]. 真菌学报, 1987, 6(3):150−152. HE S C, FENG L. A new species of the genus volvariella from Guizhou Province of China [J]. Mycosystema, 1987, 6(3): 150−152.(in Chinese)
[2]	连宾, 何绍昌, 张林, 何应华. 新型食用真菌冬小包脚菇(Volvariella brumalis)的生物学特性及营养分析 [J]. 西南农业学报, 1998, 11(1):126−128. LIAN B, HE S C, ZHANG L, et al. Biological charaeters and nutrition analysis for A newly found edible fungus, Volvariella brumalis [J]. Southwest China Journal of Agricultural Sciences, 1998, 11(1): 126−128.(in Chinese)
[3]	李瑞容, 林俊芳, 郑倩望, 等. 草菇采后生物学特性变化及保鲜技术研究进展 [J]. 中国蔬菜, 2021(6):27−33. doi: 10.19928/j.cnki.1000-6346.2021.2017 LI R R, LIN J F, ZHENG Q W, et al. Changes in biological characteristics of postharvest mushroom (Volvariella volvacea) and research progress in quality preservation technology [J]. China Vegetables, 2021(6): 27−33.(in Chinese) doi: 10.19928/j.cnki.1000-6346.2021.2017
[4]	秦惠娟, 陈屏, 王琦, 等. 草菇的化学成分、生物活性及栽培现状 [J]. 食品工业, 2017, 38(3):203−206. QIN H J, CHEN P, WANG Q, et al. The chemical composition, biological activity and cultivation situation of straw mushroom [J]. The Food Industry, 2017, 38(3): 203−206.(in Chinese)
[5]	CHEN B, WU G, LI L, et al. Effects of 1-methylcyclopropene on the quality attributes of harvested Chinese mushroom (Volvariella volvacea) fruiting bodies [J]. Food Science & Nutrition, 2019, 7(2): 747−754.
[6]	赵妍, 查磊, 陈明杰, 等. 草菇4℃贮藏期间的品质及生理生化研究 [J]. 分子植物育种, 2018, 16(21):7179−7186. doi: 10.13271/j.mpb.016.007179 ZHAO Y, ZHA L, CHEN M J, et al. Morphological and physiological indexes of straw mushroom(Volvariella volvacea) during storage at 4℃ [J]. Molecular Plant Breeding, 2018, 16(21): 7179−7186.(in Chinese) doi: 10.13271/j.mpb.016.007179
[7]	邹方伦. 贵州冬小包脚菇的生态及成分的初步研究 [J]. 贵州农业科学, 1997, 25(4):57−58. ZOU F L. Preliminary study on ecology and composition of Pleurotus eryngii in Guizhou [J]. Guizhou Agricultural Sciences, 1997, 25(4): 57−58.(in Chinese)
[8]	潘高潮, 龙汉武, 陈宵, 等. 贵州冬小包脚菇生境调查分析 [J]. 中国食用菌, 2013, 32(6):7−9. doi: 10.13629/j.cnki.53-1054.2013.06.012 PAN G C, LONG H W, CHEN X, et al. Analysis on habitat of Volvariella brumalis in Guizhou Province [J]. Edible Fungi of China, 2013, 32(6): 7−9.(in Chinese) doi: 10.13629/j.cnki.53-1054.2013.06.012
[9]	孙燕, 龙汉武, 邹方伦, 等. 冬小包脚菇形态特征及显微结构观察 [J]. 贵州科学, 2016, 34(5):22−24. doi: 10.3969/j.issn.1003-6563.2016.05.006 SUN Y, LONG H W, ZOU F L, et al. Morphological characteristics and microscopic structure of Volvariella brumalis [J]. Guizhou Science, 2016, 34(5): 22−24.(in Chinese) doi: 10.3969/j.issn.1003-6563.2016.05.006
[10]	JUSTO A, MINNIS A M, GHIGNONE S, et al. Species recognition in Pluteus and Volvopluteus (Pluteaceae, Agaricales): Morphology, geography and phylogeny [J]. Mycological Progress, 2011, 10(4): 453−479. doi: 10.1007/s11557-010-0716-z
[11]	JUSTO A, CASTRO M L. An annotated checklist of Volvariella in the Iberian Peninsula and Balearic Islands [J]. Mycotaxon, 2010, 112: 271−273. doi: 10.5248/112.271
[12]	JUSTO A, CASTRO M L. The genus Volvariella in Spain: V. dunensis comb. & stat. nov. and observations on V. earlei [J]. Mycotaxon, 2010, 112: 261−270. doi: 10.5248/112.261
[13]	徐江. 中国光柄菇属和小包脚菇属分类学研究[D]. 广州: 华南理工大学, 2016: 117. XU J. Taxonomy of Pluteus and Volvariella from china[D]. Guangzhou: South China University of Technology, 2016: 117
[14]	HIBBETT D S, BINDER M, BISCHOFF J F, et al. A higher-level phylogenetic classification of the Fungi [J]. Mycological Research., 2007, 111(5): 509−547. doi: 10.1016/j.mycres.2007.03.004
[15]	HOFFMANN K, PAWŁOWSKA J, WALTHER G, et al. The family structure of the Mucorales: A synoptic revision based on comprehensive multigene-genealogies [J]. Persoonia, 2013, 30: 57−76. doi: 10.3767/003158513X666259
[16]	SPATAFORA J W, CHANG Y, BENNY G L, et al. A Phylum-level phylogenetic classification of zygomycete fungi based on genome-scale data [J]. Mycologia, 2016, 108(5): 1028−1046. doi: 10.3852/16-042
[17]	WANI Z A, KUMAR A, SULTAN P, et al. Mortierella alpina CS10E4, an oleaginous fungal endophyte of Crocus sativus L. enhances apocarotenoid biosynthesis and stress tolerance in the host plant [J]. Scientific Reports, 2017, 7(1): 8598. doi: 10.1038/s41598-017-08974-z
[18]	JOHNSON J M, LUDWIG A, FURCH A C U, et al. The beneficial root-colonizing fungus Mortierella hyalina promotes the aerial growth of Arabidopsis and activates calcium-dependent responses that restrict Alternaria brassicae-induced disease development in roots [J]. Molecular Plant-Microbe Interactions, 2019, 32(3): 351−363. doi: 10.1094/MPMI-05-18-0115-R
[19]	HOOG G S D, GUARRO J , GENÉ J, et al. Atlas of Clinical Fungi[M]. 3rd Edition. Centraalbureau voor Schimmelcultures, Utrecht, The Netherlands/Universitat Rovira I Virgilli, Reus, Spain, 2009.
[20]	MUNDAY J, WOLFE A, LAWRENCE K, et al. Disseminated Mortierella wolfii infection in a neonatal calf [J]. New Zealand Veterinary Journal, 2010, 58(1): 62−63. doi: 10.1080/00480169.2010.65062
[21]	刘泽. 中国被孢霉属及近缘属的分类与分子系统发育研究[D]. 北京: 北京林业大学, 2020: 1-2. LIU Z. Studies on the taxonomy and molecular phylogeny of Mortierella and allied Genera in china[D]. Beijing: Beijing Forestry University, 2020: 1-2.
[22]	MARGESIN R, SCHINNER F. Properties of cold-adapted microorganisms and their potential role in biotechnology [J]. Journal of Biotechnology, 1994, 33(1): 1−14. doi: 10.1016/0168-1656(94)90093-0
[23]	FIEDUREK J, TRYTEK M, SKOWRONEK M. Strategies for improving the efficiency of bioprocesses involving toxic compounds [J]. Current Organic Chemistry, 2012, 16(24): 2946−2960. doi: 10.2174/138527212804546732
[24]	FRANCIOLI D, SCHULZ E, LENTENDU G, et al. Mineral vs. Organic amendments: Microbial community structure, activity and abundance of agriculturally relevant microbes are driven by long-term fertilization strategies. [J]. Frontiers in microbiology, 2016(7): 1446.
[25]	NING Q, CHEN L, JIA Z J, et al. Multiple long-term observations reveal a strategy for soil pH-dependent fertilization and fungal communities in support of agricultural production [J]. Agriculture Ecosystems and Environment, 2019, 293(C): 106837.
[26]	劳承英, 申章佑, 李艳英, 等. 基于高通量测序技术分析不同耕作方式下水稻根际土壤真菌多样性 [J]. 热带作物学报, 2021, 42(9):2717−2726. doi: 10.3969/j.issn.1000-2561.2021.09.038 LAO C Y, SHEN Z Y, LI Y Y, et al. Diversity analysis of fungal in rhizosphere soils of rice under different tillage methods based on high-throughput sequencing technique [J]. Chinese Journal of Tropical Crops, 2021, 42(9): 2717−2726.(in Chinese) doi: 10.3969/j.issn.1000-2561.2021.09.038
[27]	陈梦, 陈敬忠, 刘济明, 等. 小蓬竹根际土壤微生物及内生真菌多样性分析 [J]. 生态学报, 2021, 41(10):4120−4130. CHEN M, CHEN J Z, LIU J M, et al. Diversity analysis of rhizosphere soil fungi and endophytic fungi in Ampelocalamus luodianensis [J]. Acta Ecologica Sinica, 2021, 41(10): 4120−4130.(in Chinese)
[28]	张青青, 董醇波, 梁宗琦, 等. 不同产地杜仲树皮可培养内生真菌群落组成和多样性 [J]. 菌物学报, 2021, 40(10):2685−2699. doi: 10.13346/j.mycosystema.210120 ZHANG Q Q, DONG C B, LIANG Z Q, et al. Community composition and diversity of culturable endophytic fungi in bark of Eucommia ulmoides from different regions of China [J]. Mycosystema, 2021, 40(10): 2685−2699.(in Chinese) doi: 10.13346/j.mycosystema.210120
[29]	王世伟, 王卿惠, 李小鹏, 等. 镰刀菌分子鉴定与重要应用的研究进展 [J]. 微生物学通报, 2018, 45(4):907−919. doi: 10.13344/j.microbiol.china.170419 WANG S W, WANG Q H, LI X P, et al. Progress in molecular identification in the genus Fusarium and its important applications [J]. Microbiology China, 2018, 45(4): 907−919.(in Chinese) doi: 10.13344/j.microbiol.china.170419
[30]	田新莉, 蔡爱群, 曹理想, 等. 水稻内生真菌类群分析及其颉抗病原菌活性研究 [J]. 中山大学学报(自然科学版), 2005, 44(2):69−73. doi: 10.3321/j.issn:0529-6579.2005.02.018 TIAN X L, CAI A Q, CAO L X, et al. Studies on communities of endophytic fungi from rice and their antipathogenic activities in vitro [J]. Acta Scientiarum Naturalium Universitatis Sunyatseni, 2005, 44(2): 69−73.(in Chinese) doi: 10.3321/j.issn:0529-6579.2005.02.018
[31]	张清华. 油菜内生真菌的多样性及生防潜力评估[D]. 武汉: 华中农业大学, 2014. ZHANG Q H. Evaluation of the diversity and biocontrol potential of the endophytic fungi in oilseed rape[D]. Wuhan: Huazhong Agricultural University, 2014.
[32]	蔡伟建, 潘俊雅, 李济吾. 镰刀菌(Fusarium sp. )对2, 4, 6-三氯苯酚的降解特性及其机制 [J]. 环境科学学报, 2016, 36(9):3187−3192. CAI W J, PAN J Y, LI J W. Biodegradation characteristics and mechanism of 2, 4, 6-trichlorophenol by Fusarium sp. in solution [J]. Acta Scientiae Circumstantiae, 2016, 36(9): 3187−3192.(in Chinese)
[33]	谢安娜, 徐浩飞, 张志林, 等. 致病镰刀菌的研究进展 [J]. 湖北工程学院学报, 2020, 40(6):37−41. doi: 10.3969/j.issn.2095-4824.2020.06.009 XIE A N, XU H F, ZHANG Z L, et al. Research development of Fusarium [J]. Journal of Hubei Engineering University, 2020, 40(6): 37−41.(in Chinese) doi: 10.3969/j.issn.2095-4824.2020.06.009
[34]	徐雪雪, 王东, 秦舒浩, 等. 沟垄覆膜连作马铃薯根际土壤真菌多样性分析 [J]. 水土保持学报, 2015, 29(6):301−306,310. doi: 10.13870/j.cnki.stbcxb.2015.06.053 XU X X, WANG D, QIN S H, et al. Diversity analysis of fungal communities in potato continuous cropping soil under different patterns of ridge-furrow film mulching [J]. Journal of Soil and Water Conservation, 2015, 29(6): 301−306,310.(in Chinese) doi: 10.13870/j.cnki.stbcxb.2015.06.053
[35]	AGNIESZKA J, KRZYSZTOF S. Biodiversity of fungi colonising aboveground parts of zucchini ( Cucurbita pepo L. var.giromontina ) [J]. Folia Horticulturae, 2011, 23(1): 29−35. doi: 10.2478/v10245-011-0005-6
[36]	孙广正, 姚拓, 侯栋, 等. 西葫芦根腐病菌拮抗细菌的防病促生作用 [J]. 微生物学通报, 2017, 44(5):1121−1130. doi: 10.13344/j.microbiol.china.160543 SUN G Z, YAO T, HOU D, et al. Biocontrol potential of antagonistic bacteria strains against Fusarium oxysporum and their growth-promoting effects on zucchini [J]. Microbiology China, 2017, 44(5): 1121−1130.(in Chinese) doi: 10.13344/j.microbiol.china.160543
[37]	XUE C, SHEN Z, HAO Y, et al. Fumigation coupled with bio-organic fertilizer for the suppression of watermelon Fusarium wilt disease reshapes the soil microbiome [J]. Applied Soil Ecology, 2019, 140: 49−56. doi: 10.1016/j.apsoil.2019.04.007
[38]	LE H T T, PADGHAM J L, SIKORA R A. Biological control of the rice root-knot nematode Meloidogyne graminicola on rice, using endophytic and rhizosphere fungi [J]. International Journal of Pest Management, 2009, 55(1): 31−36. doi: 10.1080/09670870802450235
[39]	CARROLL G. Fungal Endophytes in Stems and Leaves: From Latent Pathogen to Mutualistic Symbiont [J]. Ecology, 1988, 69(1): 2−9. doi: 10.2307/1943154
[40]	CHUPP C. A Monograph of the Fungus Genus Cercospora [M]. Ithaca, New York: 1954.
[41]	PONS N, SUTON B C. Cercospora and similar fungi on yams (Dioscorea spp. ) [J]. Mycol Pap, 1988, 160: 1−78.
[42]	明德南. 黄瓜油壶菌和瓜类坏死斑点病毒与危地马拉甜瓜藤蔓凋萎的关系 [J]. 世界热带农业信息, 2009(2):28. MING D N. Relationship between Curcuma cucumeris and Melon Necrosis Spot Virus and the wilting of melon vines in Guatemala [J]. World Tropical Agriculture Information, 2009(2): 28.(in Chinese)
[43]	曾丽琼, 何学友, 蔡守平, 等. 白僵菌和绿僵菌感染榕管蓟马成虫试验 [J]. 防护林科技, 2019(2):8−10. doi: 10.13601/j.issn.1005-5215.2019.02.004 ZENG L Q, HE X Y, CAI S P, et al. Infection experiment of Metarhizium anisopliae and Beauveria bassiana against adults of Gynaikothrips uzeli [J]. Protection Forest Science and Technology, 2019(2): 8−10.(in Chinese) doi: 10.13601/j.issn.1005-5215.2019.02.004
[44]	蔡守平, 何学友, 曾丽琼, 等. 白僵菌和绿僵菌不同菌株对马尾松毛虫的致病力 [J]. 福建林业科技, 2018, 45(4):1−4,10. doi: 10.13428/j.cnki.fjlk.2018.04.001 CAI S P, HE X Y, ZENG L Q, et al. Comparison of pathogenicity of different Beauveria bassiana and Metarhizium anisopliae strains against Dendrolimus punctatus [J]. Journal of Fujian Forestry Science and Technology, 2018, 45(4): 1−4,10.(in Chinese) doi: 10.13428/j.cnki.fjlk.2018.04.001
[45]	INBAR J, ABRAMSKY M, COHEN D, et al. Plant growth enhancement and disease control byTrichoderma harzianum in vegetable seedlings grown under commercial conditions [J]. European Journal of Plant Pathology, 1994, 100(5): 337−346. doi: 10.1007/BF01876444
[46]	DEGENKOLB T, DIECKMANN R, NIELSEN K F, et al. The Trichoderma brevicompactum clade: A separate lineage with new species, new peptaibiotics, and mycotoxins [J]. Mycological Progress, 2008, 7(3): 177−219. doi: 10.1007/s11557-008-0563-3
[47]	MUKHERJEE P K, HORWITZ B A, KENERLEY C M. Secondary metabolism in Trichoderma: A genomic perspective[J]. Microbiology (Reading, England), 2012, 158(pt 1): 35-45.
[48]	黄毅. 食用菌栽培全一册[M]. 3版. 北京: 高等教育出版社, 2008.
[49]	黄李琳. 草菇常见病虫害及其无公害防治技术 [J]. 福建农业科技, 2017(3):38−39. doi: 10.13651/j.cnki.fjnykj.2017.03.015 HUANG L L. Common pests and diseases occurring in straw mushroom and its harmless control [J]. Fujian Agricultural Science and Technology, 2017(3): 38−39.(in Chinese) doi: 10.13651/j.cnki.fjnykj.2017.03.015