Article(id=1193476453137347519, tenantId=1146029695717560320, journalId=1190317699101192196, issueId=1193476452629836735, articleNumber=1001-2494(2025)04-0313-06, orderNo=null, doi=10.11669/cpj.2025.04.001, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1720713600000, receivedDateStr=2024-07-12, revisedDate=null, revisedDateStr=null, acceptedDate=null, acceptedDateStr=null, onlineDate=1762476560148, onlineDateStr=2025-11-07, pubDate=1740153600000, pubDateStr=2025-02-22, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1762476560148, onlineIssueDateStr=2025-11-07, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1762476560148, creator=13701087609, updateTime=1762476560148, updator=13701087609, issue=Issue{id=1193476452629836735, tenantId=1146029695717560320, journalId=1190317699101192196, year='2025', volume='60', issue='4', pageStart='313', pageEnd='438', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1762476560027, creator=13701087609, updateTime=1762482957432, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1193503285370913518, tenantId=1146029695717560320, journalId=1190317699101192196, issueId=1193476452629836735, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1193503285370913519, tenantId=1146029695717560320, journalId=1190317699101192196, issueId=1193476452629836735, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=313, endPage=318, ext={EN=ArticleExt(id=1193476453355451329, articleId=1193476453137347519, tenantId=1146029695717560320, journalId=1190317699101192196, language=EN, title=History, Current Situation and Prospect of the Armillaria associated with Gastrodia elata and Polyporus umbellatus, columnId=null, journalTitle=Chinese Pharmaceutical Journal, columnName=null, runingTitle=null, highlight=null, articleAbstract=

The symbiotic relationship between Gastrodia elata and Armillaria, as well as Polyporus umbellatus and Armillaria, is widely recognized in the field. The method of combined planting with Armillaria has emerged as a successful technique for the industrial cultivation of G. elata and P. umbellatus. However, the comprehension of the symbiotic connection between Armillaria and G. elata, as well as P. umbellatus, is still not deeply explored. The taxonomic challenge related to symbiotic Armillaria species has not received sufficient attention in scientific studies, and the incorrect utilization of Armillaria species in practical cultivation can lead to significant losses. Moreover, the limited understanding of symbiotic Armillaria groups hinders the improvement and optimal utilization of symbiotic Armillaria genetic resources. This paper conducts a retrospective analysis of the historical knowledge concerning symbiotic Armillaria with G. elata and P. umbellatus, the rationale for using Armillaria mellea, the different symbiotic Armillaria groups, and suggests potential directions for future research. The aim of this review is to enhance the understanding of symbiotic Armillaria groups, lay the groundwork for further exploration, and maximize the exploitation of symbiotic Armillaria resources.

, correspAuthors=Shunxing GUO, authorNote=null, correspAuthorsNote=null, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=null, pdfFileSize=null, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, authorCompany=null, fund=null, authors=null, authorsList=Shoujian LI, Liu LIU, Youyan LIU, Xinlei XU, Bing LI, Shunxing GUO), CN=ArticleExt(id=1193476511689830441, articleId=1193476453137347519, tenantId=1146029695717560320, journalId=1190317699101192196, language=CN, title=天麻、猪苓共生蜜环菌类群的认识历程、现状与展望, columnId=1190352408384471863, journalTitle=中国药学杂志, columnName=综述, runingTitle=null, highlight=null, articleAbstract=

天麻、猪苓与蜜环菌(Armillaria mellea)共生目前已经成为领域共识,采用蜜环菌伴栽已成为产业化生产天麻、猪苓的成功栽培模式。但研究者对于天麻和猪苓共生蜜环菌的认识仍然不够深入,在科学研究中共生蜜环菌的物种问题仍然没有引起足够重视,在实际生产中蜜环菌的错用会造成严重损失,同时共生蜜环菌类群认识的不足也限制了共生蜜环菌种质的提升与充分利用。因此,笔者梳理了天麻、猪苓共生蜜环菌类群的认识历程、分析了惯用蜜环菌的缘由、对目前天麻和猪苓共生蜜环菌的类群进行了综述,并对未来的研究方向进行了展望。以期促进对共生蜜环菌类群的认识,为共生蜜环菌的深入研究和充分利用提供依据。

, correspAuthors=郭顺星, authorNote=null, correspAuthorsNote=
*郭顺星,男,博士,教授 研究方向:药用植物内生真菌生物学 Tel:(010)57833231
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李寿建,男,博士,助理研究员 研究方向:食药用真菌遗传学与基因组学

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李寿建,男,博士,助理研究员 研究方向:食药用真菌遗传学与基因组学

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李寿建,男,博士,助理研究员 研究方向:食药用真菌遗传学与基因组学

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A-大背景下国内蜜环菌的认识历程; B-天麻共生蜜环菌认识历程;C-猪苓共生蜜环菌认识历程

, figureFileSmall=uipSQCJSJx5uvPkzH+NJ2A==, figureFileBig=i61evYJrWDL4jt16wY4hwQ==, tableContent=null), ArticleFig(id=1193540407607915471, tenantId=1146029695717560320, journalId=1190317699101192196, articleId=1193476453137347519, language=EN, label=null, caption=null, figureFileSmall=null, figureFileBig=null, tableContent=
共生物种 蜜环菌类群及发现国家
天麻Gastrodia elata A. ostoyaeA. gallicaA. sinapinaA. jezoensisA. singula,日本[55]
A. nabsnona,日本[56]
A. sinapinaA. gallicaA. mellea,中国[38]
A. gallicaA. cepistipesA. mellea,中国[39]
lineage 1(CBS L、CBS N)、lineage 2(CBS O)、lineage 3(CBS L)、lineage 6(CBS B)、A. cepistipesA. nabsnona、Nag.E,中国[17]
CBS B(lineage 6)、CBS F(A. cepistipes)、CBS N(lineage 1)、CBS K(A. mellea),中国[61]
猪苓Polyporus umbellatus A. cepistipesA. gallicaA. ostoyaeA. mellea,中国[58]
lineage 1(A. gallica)、lineage 2(A. gallica)、lineage 3(A. cepistipes)、lineage 4(A. ostoyae)、lineage 5(A. mellea),中国[59]
Clade 1(lineage 6)、Clade2、Clade 3(lineage 1)、Clade6、Clade7(lineage 4)、Clade10(A. cepistipes),中国[60]
lineage 1(CBS L、CBS N)、lineage 4(CBS J、CBS M、CBS H)、lineage 6(CBS B, Clade1、Clade2)、lineage 8(新lineage, Clade6[60])、A. cepistipes,中国[18]
), ArticleFig(id=1193540407683412944, tenantId=1146029695717560320, journalId=1190317699101192196, articleId=1193476453137347519, language=CN, label=表1, caption=

天麻、猪苓共生的蜜环菌类群及发现国家

, figureFileSmall=null, figureFileBig=null, tableContent=
共生物种 蜜环菌类群及发现国家
天麻Gastrodia elata A. ostoyaeA. gallicaA. sinapinaA. jezoensisA. singula,日本[55]
A. nabsnona,日本[56]
A. sinapinaA. gallicaA. mellea,中国[38]
A. gallicaA. cepistipesA. mellea,中国[39]
lineage 1(CBS L、CBS N)、lineage 2(CBS O)、lineage 3(CBS L)、lineage 6(CBS B)、A. cepistipesA. nabsnona、Nag.E,中国[17]
CBS B(lineage 6)、CBS F(A. cepistipes)、CBS N(lineage 1)、CBS K(A. mellea),中国[61]
猪苓Polyporus umbellatus A. cepistipesA. gallicaA. ostoyaeA. mellea,中国[58]
lineage 1(A. gallica)、lineage 2(A. gallica)、lineage 3(A. cepistipes)、lineage 4(A. ostoyae)、lineage 5(A. mellea),中国[59]
Clade 1(lineage 6)、Clade2、Clade 3(lineage 1)、Clade6、Clade7(lineage 4)、Clade10(A. cepistipes),中国[60]
lineage 1(CBS L、CBS N)、lineage 4(CBS J、CBS M、CBS H)、lineage 6(CBS B, Clade1、Clade2)、lineage 8(新lineage, Clade6[60])、A. cepistipes,中国[18]
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天麻、猪苓共生蜜环菌类群的认识历程、现状与展望
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李寿建 , 刘柳 , 刘幽言 , 许欣蕾 , 李兵 , 郭顺星 *
中国药学杂志 | 综述 2025,60(4): 313-318
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中国药学杂志 | 综述 2025, 60(4): 313-318
天麻、猪苓共生蜜环菌类群的认识历程、现状与展望
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李寿建, 刘柳, 刘幽言, 许欣蕾, 李兵, 郭顺星*
作者信息
  • 中国医学科学院北京协和医学院药用植物研究所, 北京 100193
  • 李寿建,男,博士,助理研究员 研究方向:食药用真菌遗传学与基因组学

通讯作者:

*郭顺星,男,博士,教授 研究方向:药用植物内生真菌生物学 Tel:(010)57833231
History, Current Situation and Prospect of the Armillaria associated with Gastrodia elata and Polyporus umbellatus
Shoujian LI, Liu LIU, Youyan LIU, Xinlei XU, Bing LI, Shunxing GUO*
Affiliations
  • The Institute of Medicinal Plant Development, Chinese Academy of Medical Sciences & Peking Union Medical College, Beijing 100193, China
出版时间: 2025-02-22 doi: 10.11669/cpj.2025.04.001
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天麻、猪苓与蜜环菌(Armillaria mellea)共生目前已经成为领域共识,采用蜜环菌伴栽已成为产业化生产天麻、猪苓的成功栽培模式。但研究者对于天麻和猪苓共生蜜环菌的认识仍然不够深入,在科学研究中共生蜜环菌的物种问题仍然没有引起足够重视,在实际生产中蜜环菌的错用会造成严重损失,同时共生蜜环菌类群认识的不足也限制了共生蜜环菌种质的提升与充分利用。因此,笔者梳理了天麻、猪苓共生蜜环菌类群的认识历程、分析了惯用蜜环菌的缘由、对目前天麻和猪苓共生蜜环菌的类群进行了综述,并对未来的研究方向进行了展望。以期促进对共生蜜环菌类群的认识,为共生蜜环菌的深入研究和充分利用提供依据。

蜜环菌分类  /  生物种  /  分子系统学  /  共生关系

The symbiotic relationship between Gastrodia elata and Armillaria, as well as Polyporus umbellatus and Armillaria, is widely recognized in the field. The method of combined planting with Armillaria has emerged as a successful technique for the industrial cultivation of G. elata and P. umbellatus. However, the comprehension of the symbiotic connection between Armillaria and G. elata, as well as P. umbellatus, is still not deeply explored. The taxonomic challenge related to symbiotic Armillaria species has not received sufficient attention in scientific studies, and the incorrect utilization of Armillaria species in practical cultivation can lead to significant losses. Moreover, the limited understanding of symbiotic Armillaria groups hinders the improvement and optimal utilization of symbiotic Armillaria genetic resources. This paper conducts a retrospective analysis of the historical knowledge concerning symbiotic Armillaria with G. elata and P. umbellatus, the rationale for using Armillaria mellea, the different symbiotic Armillaria groups, and suggests potential directions for future research. The aim of this review is to enhance the understanding of symbiotic Armillaria groups, lay the groundwork for further exploration, and maximize the exploitation of symbiotic Armillaria resources.

Armillaria taxonomy  /  biological species  /  molecular phylogenetic  /  symbiosis
李寿建, 刘柳, 刘幽言, 许欣蕾, 李兵, 郭顺星. 天麻、猪苓共生蜜环菌类群的认识历程、现状与展望. 中国药学杂志, 2025 , 60 (4) : 313 -318 . DOI: 10.11669/cpj.2025.04.001
Shoujian LI, Liu LIU, Youyan LIU, Xinlei XU, Bing LI, Shunxing GUO. History, Current Situation and Prospect of the Armillaria associated with Gastrodia elata and Polyporus umbellatus[J]. Chinese Pharmaceutical Journal, 2025 , 60 (4) : 313 -318 . DOI: 10.11669/cpj.2025.04.001
蜜环菌是蜜环菌属[Armillaria(Fr.) Staude]真菌的统称,目前全世界已定名的蜜环菌生物种有近40种[1],国内已报道的蜜环菌生物种有16种,其中包含已鉴定的蜜环菌7种,分别为Armillaria gallicaA. ostoyaeA. melleaA. sinapinaA. borealisA. korhonenii[2]和目前已经转移至Desarmillaria属的D. tabescens[3],未鉴定的9种,分别为中国生物种(Chinese biological species,CBS)C、CBS F、CBS G、CBS H、CBS J、CBS L、CBS N、CBS O和CBS P[4-9]
蜜环菌广泛分布于温带到热带地区,生态类型复杂,其既可以营腐生生活、寄生生活[10],又可以与兰科植物天麻(Gastrodia elata)和多孔菌科真菌猪苓(Polyporus umbellatus)共生[11-12]。天麻和猪苓是我国传统中药,天麻具有熄风止痉、平抑肝阳、祛风通络、补益之效,常用来治疗眩晕眼黑、头风头痛、肢体麻木、半身不遂、风湿痹痛及惊痫等症[13],猪苓具有利水渗湿的功效,用于水肿、小便不利、泄泻、淋浊、带下等病症[14]。虽然两者分属于植物界和真菌界,但两者均能与蜜环菌Armillaria形成共生关系。天麻、猪苓和蜜环菌的共生关系可以概括为“非你不可”,即脱离了蜜环菌则天麻和猪苓无法单独生长。而二十世纪六七十年代国内学者对于天麻、猪苓与蜜环菌共生关系的发现和认识,也使得我国率先解决了天麻和猪苓无法人工栽培的难题[11,15]
天麻与蜜环菌共生关系的报道可追溯至1911年[16],猪苓与蜜环菌共生关系的报道可追溯至1978年[15]。随着生物种鉴定方法的建立和分子系统学的发展,蜜环菌的分类体系逐渐完善,国内蜜环菌的类群也逐渐清晰[2],天麻、猪苓共生的蜜环菌类群得以揭示[17-18]。但目前关于天麻、猪苓的研究中仍广泛记载天麻、猪苓与蜜环菌(A. mellea)共生,已严重落后于现今对于共生蜜环菌的认识,因此本研究旨在综述天麻、猪苓共生蜜环菌的认识历程、惯用A. mellea的缘由以及共生蜜环菌类群的研究现状,同时对未来的研究方向进行了展望,以期引起天麻、猪苓研究者的重视,促进天麻、猪苓共生蜜环菌资源的充分利用和优良菌株的选育,进而促进天麻、猪苓产业的持续健康发展。
A. mellea隶属于蜜环菌属,是蜜环菌属众多物种中的其中一种。自1911年首次记载天麻与蜜环菌A. mellea共生[16],1981年首次记载猪苓与蜜环菌A. mellea共生[19],目前该名称已在文献中广泛记载与天麻和猪苓共生[20-25]。逐年查阅目前国内发表的关于天麻、猪苓的文章,发现几乎有记载的年份中均出现共生蜜环菌为A. mellea的记载。即使目前已经有学者提出天麻、猪苓共生蜜环菌为多个蜜环菌物种的情况下,A. mellea仍然广泛使用[26-29],其中有相关研究在文中进行了分子序列的比对,得出结果为与A. gallica更为相似,但仍然在题目中使用了A. mellea这一名称[28]
天麻与蜜环菌的共生关系认识较早,自1911年日本学者Kusano首次报道天麻与蜜环菌共生则使用了A. mellea[16],而后国内学者周铉1973年首次在国内描述天麻与蜜环菌的共生关系便沿用了该名称[30],此后被称为“天麻之父”的徐锦堂和周铉在研究中也均沿用了这一名称[31-37]。自1973年至今有大量学者在天麻相关的研究论文中仍采用该名称[22-26,28-29],即使在2008年后开始有学者证明了天麻共生的蜜环菌包含了多个蜜环菌属的物种[38-39],这一学名仍然广泛使用。
相对于天麻,猪苓与蜜环菌共生关系的记载较晚,自1978年蜜环菌伴栽猪苓栽培成功后[15],1981年李世全首次在文献中记载与猪苓共生的蜜环菌学名为Armillariella mellea[19],此后1991年徐锦堂记载为Armillaria mellea[40],在1991—1996年间,两个名称混杂使用[41-44],1996年后均记载为A. mellea,至今这一名称也仍然在猪苓相关研究中使用[20,27]。由于猪苓与天麻栽培方式几乎相同,同时猪苓与蜜环菌的共生关系在天麻与蜜环菌共生关系的认识基础上发现,因此猪苓共生蜜环菌学名的使用可能受当时天麻研究的影响,也沿用了这一学名。因此历史沿用可能是共生蜜环菌惯用A. mellea的重要原因。
自林奈提出“双名法”之后,各种生物都开始使用拉丁学名作为其通用学名。对于国内报道的物种,一般会对应有中文名,蜜环菌属物种也不例外。国内已经鉴定的蜜环菌物种有高卢蜜环菌、奥氏蜜环菌、芥黄蜜环菌、北方蜜环菌、假蜜环菌、科赫宁蜜环菌和蜜环菌,其对应的拉丁学名分别为A. gallicaA. ostoyaeA. sinapinaA. borealisA. tabesecensA. korhoneniiA. mellea[4-6,8,45]。由此可以看出,不同于其他蜜环菌,A. mellea的中文名即为“蜜环菌”,大家常说天麻、猪苓与蜜环菌共生,而以蜜环菌作为名称对应的拉丁学名则为A. mellea。因此容易理解为蜜环菌与天麻、猪苓共生就是A. mellea与天麻、猪苓共生,这可能是惯用A. mellea的另一个重要原因。
由于小部分文献记载中也使用了Armillariella mellea一名,因此对此进行解释。Armillariella属与Armillaria属是同物异名,Armillaria属于1857年被Staude提出,Armillariella属于1881年被Karsten提出[45],并且两个属使用了相同的模式种,根据国际植物命名法规(International Code of Botanical Nomenclature)第14.4条,ArmillariellaArmillaria的晚出异名,因此Armillaria应为该属正确的学名[45-46]
天麻、猪苓共生蜜环菌的认识是在蜜环菌认识的大背景下进行的。蜜环菌属最初在1821年作为蘑菇属Agaricus下的一个族(tribe)提出,至1857年Staude将其提升为蜜环菌属Armillaria(图1A),虽然后续对于是谁最早合法提出Armillaria属有过争议,比如Singer在不同版本的“Agaricales in Modern Taxonomy”中分别记载了认为Quélet(1872)和Kummer(1871)作为蜜环菌属的首次合法提出人,认为Staude只是将Armillaria提升到了属水平,但在该属内未描述任何新物种或新组合。后续根据对Kummer文献的分析以及Kummer提出蜜环菌属时模式物种使用的错误,最后大家广泛接受Staude(1857)的描述满足了所有当时合格发表的要求,Armillaria Staude是有效的合法名称[45-46]
此后蜜环菌分类的又一进步是1973年Hintikka发现了蜜环菌的四极性交配系统[47],而后Korhonen(1978)和Anderson & Ullrich(1979)利用单倍体菌株交配鉴定了欧洲和美洲的蜜环菌生物种(biological species)[48-49],此后通过单倍体交配判定生物种成为了蜜环菌主要的分类方法[50](图1A)。在世界蜜环菌生物种研究的热潮中,国内也于20世纪90年代开始了蜜环菌生物种的研究,1996年He等[4]首先发表了国内大兴安岭和长白山地区的5个蜜环菌生物种CBS A、CBS B、CBS C、CBS D、CBS E,并通过与欧洲蜜环菌生物种进行交配验证,证明其中两种为高卢蜜环菌A. gallica(CBS B)和奥氏蜜环菌A. ostoyae(CBS E)。此后秦国夫和赵俊等在国内不断进行生物种研究,1999年Qin等[5]证明中国生物种CBS A为芥黄蜜环菌A. sinapina,中国生物种CBS D也为A. ostoyae。2000年Qin等[6]进一步发现了4个蜜环菌新生物种CBS F、CBS G、CBS H、CBS I,其中CBS I为假蜜环菌A. tabescens(Desarmillaria tabescens)。2005年,Zhao等[8]发现了6个蜜环菌新生物种,CBS J、CBS K、CBS L、CBS M、CBS N、CBS O,其中CBS K为A. mellea,CBS M为A. borealis。2008年,Zhao等[9]发现了蜜环菌中国新生物种CBS P(图1A)。至此中国蜜环菌生物种的研究已经基本结束,直至2024年Liu等[2]再次发现了中国蜜环菌新生物种CBS Q,并命名为A. korhonenii。目前中国已报道的蜜环菌生物种共计16种(图1A)。
对于天麻共生的蜜环菌,1911年由日本学者Kusano首次发现[16](图1B),当时蜜环菌已提升至蜜环菌属,并且该属的模式物种为Armillaria mellea(1871, https://www.species fungorum.org/Names/SynSpecies.asp?RecordID=190066)。至1973年,周铉首次在国内文献记载天麻与蜜环菌共生,沿用了A. mellea这一名称[30],此后基于蜜环菌生物种研究背景,秦国夫于2005年登记科技成果“蜜环菌优良高产杂交菌株栽培天麻技术”,当时对于蜜环菌生物种已经有了部分认识,但公开的登记信息中并未描述所用蜜环菌物种。2007年Zhao等[51]在文献中记载不同种类的天麻可能需要不同的蜜环菌类群,揭开了天麻共生蜜环菌类群研究的序幕,此后2008年Ji[52]鉴定天麻共生蜜环菌为高卢蜜环菌,2011年彭述敏通过ITS(internal transcribed spacer,内转录间隔区)鉴定天麻共生蜜环菌与A. sinapinaA. gallicaA. mellea同源性最高[38],2014年Huang等[39]通过ITS鉴定天麻共生蜜环菌属于A. gallicaA. cepistipesA. mellea。2016年Guo等[17]基于ITS、tef-1α(translation elongation factor-1α,翻译延长因子1α)和β-tubulin(β微管蛋白)序列进行分子系统学研究,发现天麻共生蜜环菌分布在7个系统发育谱系(图1B),此后开始有学者基于该结果进行天麻共生蜜环菌的物种鉴定[53-54]。日本学者在1995年报道了天麻共生蜜环菌5种,分别为A. ostoyaeA. gallicaA. sinapinaA. jezoensisA. singula,2008年报道了天麻共生蜜环菌1种,为A. nabsnona[55-56],至此天麻共生蜜环菌的认识已较为清晰。
对于猪苓共生蜜环菌的认识,除了在蜜环菌认识历程下进行之外,也是在天麻共生蜜环菌的认识基础进行的(图1C)。1978年山西古县猪苓场首次报道蜜环菌伴栽猪苓成功[15],此后1980年Zhang等[57]初步探讨猪苓和蜜环菌的关系。1981年Li等[19]首次在文献中记载猪苓与蜜环菌Armillariella mellea共生,此后1991年Xu等[40]记载猪苓共生蜜环菌为Armillaria mellea,一直到1996年2个名称处于混用状态[41-44],1996年之后基本再没有Armillariella mellea的使用。直至2015年Liu等[58-59]基于IGS(intergenic spacer,转录间隔区)序列先后报道了猪苓共生的蜜环菌为4个种、5个系统发育谱系。此后Men等[60]和Xing等[18]2017年结合ITS、tef-1αβ-tubulin片段进一步报道了猪苓共生蜜环菌分布在5个系统发育谱系[18](图1C),至此猪苓共生蜜环菌的认识已较为清晰。
目前业界对于天麻和猪苓共生的蜜环菌类群有了较为系统的了解。对于天麻共生的蜜环菌,此前日本学者记载在日本A. ostoyaeA. gallicaA. sinapinaA. jezoensisA. singulaA. nabsnona能与天麻共生[55-56]。国内Jin[52]、Peng[38]和Huang等[39]也对天麻共生蜜环菌进行了鉴定,直至2016年Guo等[17]对国内天麻共生的蜜环菌做了较为系统的工作,发现国内天麻共生蜜环菌聚类到7个系统发育谱系。此后,Yang等[61]记载进行天麻栽培的蜜环菌材料有4个蜜环菌生物种(表1)。总结国内记载与天麻共生的蜜环菌类群,包括已经定名的6种,包括A. sinapinaA. gallicaA. melleaA. cepistipesA. nabsnona和Nag. E,未定名的生物种 lineage 1(CBS L、CBS N)、CBS O(lineage 2)、CBS L(lineage 3)、CBS B(lineage 6)。
对于猪苓共生的蜜环菌,2010年日本学者Kikuchi和Yamaji基于ITS序列进行了猪苓共生蜜环菌的鉴定,但由于系统发育树上不能完全分开,因此未鉴定到具体的物种[62]。此后,2015年Liu等[58-59]对猪苓共生蜜环菌类群进行研究,2017年Men等[60]和Xing等[18]又做了进一步的研究,基本摸清了猪苓共生的蜜环菌类群。总结记载与猪苓共生的蜜环菌类群,包括已定名的A. cepistipesA. gallicaA. ostoyaeA. mellea,未定名的CBS B、lineage1(CBS L、CBS N)、lineage4(CBS J、CBS M、CBS H)和新发现的lineage8。
目前对于天麻、猪苓共生的蜜环菌类群已经有了基本的认识,明确了天麻、猪苓共生的蜜环菌类群并非单一的蜜环菌物种,而是包含了多种不同的蜜环菌。但这些研究至今仅有十余年的时间,研究范围也具有一定的局限性。Guo等[17]采集的天麻共生蜜环菌菌株大多集中在云南地区,Yang等[61]采集的天麻共生蜜环菌菌株在鄂西南地区,Ji[52]仅在吉林部分地区进行了天麻共生蜜环菌采集,Peng[38]和Huang等[39]也仅在云南、贵州地区收集了天麻共生蜜环菌,总体来看天麻共生蜜环菌的资源收集范围不够广泛。对于猪苓共生蜜环菌,资源较为广泛,遍布黑龙江、吉林、河北、河南、陕西、云南等多个省份[18,58-60],但从材料数量上来看仍然具有较大的提高空间。因此未来更大覆盖范围内资源的收集,将会更为全面地揭示天麻、猪苓共生的蜜环菌类群。
目前人工栽培的天麻主要为乌天麻Gastrodia elata f.glauca、红天麻Gastrodia elata f. elata和绿天麻Gastrodia elata f. viridis。Guo等[17]在研究共生蜜环菌类群时区分了红天麻与乌天麻,根据聚类结果反映红天麻共生蜜环菌主要集中在lineage 6,乌天麻共生蜜环菌分散在多个lineage中,而杨笑等记载的红天麻共生蜜环菌类群对应到Guo等[61]的分子系统树中,也分散在多个系统发育谱系中。目前对于不同的天麻变形对应的蜜环菌类群是否有差别,是否有针对不同天麻有专一性的蜜环菌类群还不清楚。猪苓为单一物种,但不同地理分布的猪苓遗传差异较大[63],Xing等[18]通过系统发育网络分析,结果显示亲缘关系较近的猪苓倾向于与亲缘关系较近的蜜环菌类群进行共生。那么相同的天麻变型亲缘关系的远近是否也对蜜环菌类群的选择具有倾向性还缺乏相关研究。此外,不同地区之间是否有本地优势的共生蜜环菌类群仍需要进一步揭示,这些研究将对共生蜜环菌的选择和利用具有重要意义。
用于天麻、猪苓栽培的蜜环菌菌株主要分离于天麻块茎和猪苓菌核上的菌索,而自然界中独立于天麻、猪苓的蜜环菌菌索和子实体资源基本没有利用。虽然共生需要地理位置上的接近,但独立于天麻、猪苓的蜜环菌也未必不能与天麻、猪苓共生。蜜环菌广泛分布在温带到热带地区,国内分布了大量的蜜环菌资源,据Qin等[7]采集的蜜环菌标本记载国内有15个省份以上有蜜环菌分布,综合对于各地蜜环菌的记载国内有20个以上的省份分布有蜜环菌资源[9,64],按目前鉴定的天麻、猪苓共生的蜜环菌类群,很多野生采集的蜜环菌子实体与共生蜜环菌属于相同物种,同时产区也存在生产者将采自山上分离的蜜环菌种植天麻、猪苓的现象,这也说明这些未在天麻、猪苓上发现的蜜环菌资源可能是未来优质共生蜜环菌的重要来源。
人为选种只能在自然资源中筛选菌株,无法进行菌株的创制和提升。目前天麻和猪苓在全国栽培范围广、生产气候差异大,因此对蜜环菌的要求较高,盲目引种会造成生产失败,而目前缺乏普适性的蜜环菌生产菌株。因此除充分利用在天麻、猪苓上分离的蜜环菌之外的蜜环菌资源,还可以通过育种手段针对生产环境进行蜜环菌种质的创制和提升。共生蜜环菌一般通过菌索分离,得到的是其双倍体菌株,进行伴栽筛选,无法人为进行种质提升。在蜜环菌生物种广泛验证的背景下,进行杂交育种相对容易实现。野外形成子实体的蜜环菌易于获得其单倍体菌株,可以通过杂交手段进行蜜环菌种质的提升。同时文献记载,目前有些蜜环菌类群如奥氏蜜环菌A. ostoyae、蜜环菌A. mellea、假蜜环菌A. tabescens、高卢蜜环菌A. gallica可以通过人工诱导形成子实体[45,65],本实验室前期也成功诱导获得了猪苓共生的奥氏蜜环菌的子实体,因此未来可以通过杂交育种手段提升共生蜜环菌种质,从而促进天麻和猪苓产业的发展。
  • 北京协和医学院中央高校基本科研业务费资助(3332023052)
  • 中国医学科学院医学与健康科技创新工程资助(2021-I2M-1-031)
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doi: 10.11669/cpj.2025.04.001
  • 接收时间:2024-07-12
  • 首发时间:2025-11-07
  • 出版时间:2025-02-22
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  • 收稿日期:2024-07-12
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北京协和医学院中央高校基本科研业务费资助(3332023052)
中国医学科学院医学与健康科技创新工程资助(2021-I2M-1-031)
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    中国医学科学院北京协和医学院药用植物研究所, 北京 100193

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*郭顺星,男,博士,教授 研究方向:药用植物内生真菌生物学 Tel:(010)57833231
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2种不同金属材料的力学参数

Family
属数
Number of
genus
种数
Number of
species
占总种数比例
Percentage of
total species (%)

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