微生物学报

Fruiting body types	Dominant strain (phylum/genus)	Analytic method	References
Black truffle	Proteobacteria Bacteroidetes	High-throughput sequencing	[38]
Cantharellus spp.	Hafnia Stenotrophomoas Pseudomonas	BOX-PCR fingerprinting	[43]
C. cibarius	Proteobacteria Bacteroidetes	High-throughput sequencing	[42]
Tricholoma matsutake	Proteobacteria Bacteroidetes Firmicutes	High-throughput sequencing	[26]
T. matsutake	Micrococcales Bacillales Caulobacter	High-throughput sequencing	[44]
Sanghuangporus	Proteobacteria Bacteroidota Firmicutes	High-throughput sequencing	[45]
T. matsutake	Proteobacteria Firmicutes	Culture-dependent method	[46]
T. ganbajun	Pseudomonas	Culture-dependent method	[47]
Suillus grevillei	Pseudomonas Bacillus	Culture-dependent method	[48]
T. aestivum	Pseudomonas Buttiauxella	Culture-dependent method	[15]

Fruiting body types	Dominant strain (phylum/genus)	Analytic method	References
Black truffle	Proteobacteria Bacteroidetes	High-throughput sequencing	[38]
Cantharellus spp.	Hafnia Stenotrophomoas Pseudomonas	BOX-PCR fingerprinting	[43]
C. cibarius	Proteobacteria Bacteroidetes	High-throughput sequencing	[42]
Tricholoma matsutake	Proteobacteria Bacteroidetes Firmicutes	High-throughput sequencing	[26]
T. matsutake	Micrococcales Bacillales Caulobacter	High-throughput sequencing	[44]
Sanghuangporus	Proteobacteria Bacteroidota Firmicutes	High-throughput sequencing	[45]
T. matsutake	Proteobacteria Firmicutes	Culture-dependent method	[46]
T. ganbajun	Pseudomonas	Culture-dependent method	[47]
Suillus grevillei	Pseudomonas Bacillus	Culture-dependent method	[48]
T. aestivum	Pseudomonas Buttiauxella	Culture-dependent method	[15]

Host fungi	Associated microbes	Activities	References
Agaricus bisporus	Bacillus	Anti-bacteria/fungi	[16]
Tuber borchii	Actinomycete Pseudomonad	Anti-fungi	[59]
Tricholoma matsutake	Ewingella americana Pseudomonas endophytica Serratia marcescens Mycetocola lacteus Cedecea neteri	Anti-fungi	[45]
Volvariella volvacea	Bacillus sp. C2.2, C3.8, D3.3	Antifungal activities against Cercospora lactucae, C. gloeosporioides, Fusarium oxysporum	[60]
A. bisporus	Pseudomonas sp. AB-114	Anti-bacteria/fungi	[61]
T. matsutake	Stenotrophomonas maltophilia	Chitinase, endoglucanase, β-glucosidase, protease, lipase activities	[45]
T. borchii	Pseudomonas	Indole-3-acetic acid, siderophores, protease, collagenase, carboxymethyl cellulose activities	[62]
Shiraia sp. S9	P. fulva SB1	Antioxidation	[63]
Inonotus obliquus	Acremonium sp.	Antitumor	[64]

Host fungi	Associated microbes	Activities	References
Agaricus bisporus	Bacillus	Anti-bacteria/fungi	[16]
Tuber borchii	Actinomycete Pseudomonad	Anti-fungi	[59]
Tricholoma matsutake	Ewingella americana Pseudomonas endophytica Serratia marcescens Mycetocola lacteus Cedecea neteri	Anti-fungi	[45]
Volvariella volvacea	Bacillus sp. C2.2, C3.8, D3.3	Antifungal activities against Cercospora lactucae, C. gloeosporioides, Fusarium oxysporum	[60]
A. bisporus	Pseudomonas sp. AB-114	Anti-bacteria/fungi	[61]
T. matsutake	Stenotrophomonas maltophilia	Chitinase, endoglucanase, β-glucosidase, protease, lipase activities	[45]
T. borchii	Pseudomonas	Indole-3-acetic acid, siderophores, protease, collagenase, carboxymethyl cellulose activities	[62]
Shiraia sp. S9	P. fulva SB1	Antioxidation	[63]
Inonotus obliquus	Acremonium sp.	Antitumor	[64]

子实体伴生菌多样性及其生物活性研究进展

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沈文浩 ¹ , 郑丽屏 ² , 周建芹 ¹ , 王剑文 ¹^,^*

微生物学报 | 微生物资源多样性 2025,65(4): 1433-1445

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微生物学报 | 微生物资源多样性 2025, 65(4): 1433-1445

子实体伴生菌多样性及其生物活性研究进展

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沈文浩¹, 郑丽屏², 周建芹¹, 王剑文¹^,^*

作者信息

^1.苏州大学药学院，江苏苏州

^2.苏州大学金螳螂建筑学院，江苏苏州

Research progress in the diversity and biological activities of fruiting body-associated microbes

Wenhao SHEN¹, Liping ZHENG², Jianqin ZHOU¹, Jianwen WANG¹^,^*

Affiliations

^1.College of Pharmaceutical Sciences, Soochow University, Suzhou, Jiangsu, China

^2.Gold Mantis School of Architecture, Soochow University, Suzhou, Jiangsu, China

出版时间: 2025-04-04 doi: 10.13343/j.cnki.wsxb.20250010

文章导航

摘要

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真菌子实体是重要的药物和食物资源，其中存在丰富多样的伴生微生物。目前，关于子实体内部细菌群落组成的报道已陆续发表，但对于真菌子实体伴生菌的多样性及其生物活性仍知之甚少。子实体与伴生菌之间形成了一个赖以生存的生命有机体，伴生菌能够促进宿主真菌生长，增强其适应性，提高对环境胁迫的抗性，并促进其次生代谢产物的积累。此外，子实体伴生菌还具有抗菌、抗氧化以及抗肿瘤等生物医药活性。本文综述了近年来子实体伴生菌的分离与培养方法，分析了伴生细菌、放线菌和真菌群落多样性，进一步挖掘了子实体伴生菌的生物活性。本文为伴生菌与寄主关系的研究提供了参考，并为子实体伴生菌资源的开发利用奠定了基础。

关键词

子实体 / 伴生菌 / 多样性 / 真菌发育 / 次生代谢

Abstract

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Fruiting bodies represent pivotal resources with medicinal and edible values, hosting a diverse array of microbes. Although studies have elucidated the composition and structures of bacterial communities inhabiting fruiting bodies, the diversity and biological functions of fruiting body-associated microbes remains elusive. The symbiotic associations between fruiting bodies and their microbial inhabitants play integral roles in promoting the growth, enhancing the adaptation to environmental stresses, and facilitating the accumulation of secondary metabolites of host fungi. Additionally, these fruiting body-associated microbes exhibit promising biomedical properties, including antibacterial, antioxidant, and anti-tumor activities. This paper reviews the recent advancements in the isolation and cultivation techniques of these associated microbes, delineates the diversity of associated bacteria, actinomycetes, and fungi, elucidates their biological activities, and provide insights into the intricate interactions between associated microbes and their host fungi. This review offers avenues for the future research and utilization of fruiting body-associated microbes.

Key words

fruiting bodies / associated microbes / diversity / fungal development / secondary metabolism

引用本文

沈文浩, 郑丽屏, 周建芹, 王剑文. 子实体伴生菌多样性及其生物活性研究进展. 微生物学报, 2025 , 65 (4) : 1433 -1445 . DOI: 10.13343/j.cnki.wsxb.20250010

Wenhao SHEN, Liping ZHENG, Jianqin ZHOU, Jianwen WANG. Research progress in the diversity and biological activities of fruiting body-associated microbes[J]. Acta Microbiologica Sinica, 2025 , 65 (4) : 1433 -1445 . DOI: 10.13343/j.cnki.wsxb.20250010

正文

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真菌子实体(fruiting bodies)是真菌在其生命周期中形成的多细胞结构，主要功能是产生并传播有性孢子(如担孢子或子囊孢子)，从而实现繁殖，是真菌从营养生长(菌丝体阶段)转向生殖生长的重要标志^[1]。一些真菌子实体是天然绿色食材和药源宝库，其代谢产物结构多样、新颖且独特，在抗肿瘤、提高免疫力、延缓衰老等方面具有较强的生理活性。在食用菌栽培过程中，真菌子实体的侵染性病原已屡见不鲜，但在子实体生长过程中，一类非致病的微生物也常常伴随子实体一起生长和发育，它们定殖于宿主真菌的孢子、菌丝体表面以及细胞内部，然而目前尚未有一个准确的概念来定义这类微生物。目前，大部分学者用子实体“伴生菌”^[2-4]来描述这类微生物，《食用菌术语(GB/T 12728—2006)》^[5]将“伴生现象”定义为“2种真菌共同生存在同一基物上，其中一种对另外一种的生长发育有促进作用”。也有一些学者分别用“内生菌(endophytes)”^[6]和“附生菌(epiphytic bacteria)”^[7]来定义这类从子实体内分离的微生物。内生菌一般特指定殖于健康植物组织内部的微生物，而附生菌常用于描述附着于生物体表面且不深入组织内部的细菌。子实体伴生菌(fruiting body-associated microbes)与宿主菌的关系根据环境条件及相互作用不同而改变，可能是相互有益、中性的或具有潜在危害性^[8]。辛智海^[9]报道了采用纯培养法从鸡油菌(Cantharellus cibarius)子实体中分离的一株伴生细菌对鸡油菌的孢子萌发有显著促进作用。李辉等^[4]通过变性梯度凝胶电泳(denatured gradient gel electrophoresis, DGGE)技术研究了来自8种不同地区的冬虫夏草(Cordyceps sinensis)子实体中的细菌群落结构，其中以变形菌门(Proteobacteria，50.0%)、拟杆菌门(Bacteroidetes，33.0%)和厚壁菌门(Firmicutes，16.0%)为优势菌门。李强等^[6]采用纯培养法从翘鳞肉齿菌[Sarcodon imbricatus (L.ex Fr.) Karst.]子实体中分离出52株内生细菌和26株内生真菌，细菌以伯克氏菌属(Burkholderia，40.4%)和假单胞菌属(Pseudomonas，23.1%)为优势菌属，真菌以Phaeomoniella (26.9%)和隐球菌属(Cryptococcus，11.5%)为优势菌属。张瑞颖等^[7]对从白灵侧耳(Pleurotus nebrodensis)子实体中分离的75株附生细菌进行了鉴定，以节杆菌属(Arthrobacter)和假单胞菌属(Pseudomonas)为优势属群。目前，对于子实体伴生菌的多样性及其生物功能的研究尚无综述文章。本文介绍了子实体伴生菌分离与培养的方法，归纳了不同类型子实体中伴生菌群落的多样性，并探讨了子实体伴生菌的生物功能，旨在为真菌子实体与伴生菌的互作关系研究提供基础，并为伴生菌的生物医药潜能利用提供参考。

1 子实体伴生菌多样性分析

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近年来，植物或子实体内部微生物群落的生物特性、多样性及外界因素导致的种群动态变化，以及这些微生物产生的新颖活性代谢产物受到了极大关注。目前，微生物群落多样性的主要分析方法包括纯培养法(culture-dependent method)与非培养法(culture-independent method)^[10]。这2种方法是研究子实体内部微生物群落结构和挖掘其中微生物资源的基础。近年来，非培养法，如高通量测序法(high-throughput sequencing)，也被用于检测整体微生物群落的变化^[11-12]。

1.1 纯培养法

纯培养法(culture-dependent method)是最早用于计量微生物群落结构的方法，可以检测环境条件改变导致的微生物群落变化^[13]。纯培养法通过分离培养基从样本中分离出目标微生物菌株，通过观察微生物的外形(如菌丝形态)、生理生化特性，并结合分子生物学手段对分离菌株的16S rRNA基因进行鉴定。对于样本内的微生物(如植物内生菌或子实体伴生菌)的分离，需要对样本进行预处理和表面充分消毒杀菌后，再转移到相应的培养基上分离。表面灭菌的主要方法分为物理灭菌(包括紫外杀菌、火焰灭菌)^[14]和化学灭菌^[6]。化学灭菌较为常见，一般使用表面消毒剂，如表面活性剂、次氯酸钠、硫代硫酸钠、乙醇和升汞等进行灭菌。李云龙等^[15]采用化学灭菌法从多株夏块菌(Tuber aestivum)子实体中分离出233株细菌，属于5门11属。

从特定植物组织中分离出的内生菌的数量和多样性可能会因生长培养基的选择而有所不同，尚无一种培养基可以满足所有细菌的营养和生长需求^[16]。因此，针对不同的分离菌株，需使用专门的培养基对样本特定位置的特定微生物群落进行分离和纯化。使用马铃薯葡萄糖琼脂(potato dextrose agar, PDA)培养基分离样本内的真菌；细菌的分离常采用LB固体培养基；放线菌分离的培养基种类相对较多。Xiang等^[17]采用牛肉-蛋白胨分离培养基从双孢蘑菇(Agaricus bisporus)子实体中分离出55株细菌和放线菌，其中以放线菌为优势种群。张兆娟等^[18]采用高氏一号培养基从网柄菌(dictyostelids)子实体中分离出28株放线菌，其中以微杆菌属(Microbacterium)为优势种群。

1.2 非培养法

非培养法是直接从环境样本中提取DNA，进行16S rRNA基因的PCR扩增，并结合系统发育分析，这种方法揭示了大量以前未知和难培养的微生物。在一个微生物多样性丰富的环境中，纯培养法只能分离出符合生长需求条件的细菌，但大多数内生菌的生长条件处于未知状态，一些依赖于特殊环境或宿主资源的微生物在实验条件下无法生长。在实验室条件下，只有不到1%的细菌种类是可培养的^[19]，表明如果仅采用纯培养法来分析样本的多样性，绝大多数微生物群落可能不会被检测到^[19-20]。截至目前，已报道微生物可以产生2万多种生物活性物质^[21]，这也意味着大量的微生物资源尚未被挖掘。近年来，研究开发了多种用于检测微生物多样性的非培养法。大部分非培养方法都是基于聚合酶链反应(polymerase chain reaction, PCR)技术，如变性/温度梯度凝胶电泳(denaturing/temperature gradient gel electrophoresis, DGGE/TGGE)^[22]、单链构象多态性(single strand-conformation polymorphism, SSCP)^[23]、限制片段长度多态性(restriction fragment length polymorphisms, RFLP)^[24]、末端限制性片段长度多态性(terminal restriction fragment length polymorphisms, T-RFLP)^[25]、焦磷酸测序(barcoded pyrosequencing)^[26]、定量聚合酶链反应(quantitative PCR, qPCR)^[27]和高通量测序法(high-throughput sequencing)^[28]。其他不依赖PCR等分子技术的方法，如微阵列(microarray)和荧光原位杂交(fluorescence in situ hybridization, FISH)，也已被用于微生物多样性分析^[29-30]。一些基于单细胞的技术，如拉曼显微光谱(Raman microspectroscopy)和纳米级二级离子质谱(nano-scale secondary ion mass spectrometry, NanoSIMS)，也逐渐应用于微生物生态学和环境微生物学领域^[31-32]。其中，高通量测序技术凭借其经济、高效的特点被广泛用于微生物群落多样性的研究，它能够对样品中优势群落、稀有菌群、难培养的菌群及未知菌种进行检测。Li等^[26]通过利用焦磷酸测序(barcoded pyrosequencing)发现，来自四川多地区的松茸(Tricholoma matsutake)子实体中以厚壁菌门(Firmicutes)、变形菌门(Proteobacteria)及拟杆菌门(Bacteroidetes)的伴生菌为主。Benucci等^[33]通过高通量测序(high-throughput sequencing)比较了羊肚菌(Morchella sextelata)栽培土壤中不同部位(毛、茎、土壤)的原核生物群落，发现整个细菌群落主要以拟杆菌门(Bacteroidota，36.7%)和变形菌门(Proteobacteria，23.7%)为主。Chen等^[34]利用高通量测序技术发现，不同地区的黑松露(Tuber indicum)子实体中以根瘤菌(Bradyrhizobium)为优势属。其中，微生物受地理位置和土壤特征的影响，子实体中的细菌群落的丰度和多样性高于外生菌根和土壤中的细菌群落。通过非培养法还可以对微生物群落的功能作用进行检测。Liu等^[35]采用功能基因微阵列(microarray)和扩增测序技术分析干巴菌(Thelephora ganbajun)子实体内部的微生物多样性及其功能基因，发现其中1 334个基因参与碳和氮的固定，524个基因影响中部子实层的碳和亚硫酸盐还原，255个基因参与菌盖皮层的硫氧化和多磷酸盐降解。Ren等^[36]通过高通量测序发现，在灵芝(Ganoderma lucidum)培养前后土壤中的微生物群落存在显著差异，子囊菌门(Ascomycota)、担子菌门(Basidiomycota)、变形菌门和放线菌门在灵芝培养后表现出明显差异，且真菌和细菌群落结构与土壤pH、氮、碳和微量元素(Fe、Zn、Mn、Mn、Cu)含量显著相关。

2 子实体伴生菌的分类

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2.1 子实体伴生细菌

细菌在子实体微生物群落中占比较大，如在松露的内部和外部，细菌密集定殖，其密度可达每克子实体(干重) 10⁸个细胞^[37]。Antony‐Babu等^[38]通过高通量测序技术分析了黑松露(T. melanosporum)微生物群落的多样性，发现变形菌门(Proteobacteria，11%-21%)和拟杆菌门(Bacteroidetes，2%-20%)为优势种群。大部分松露被复杂的细菌群落定殖，主要由变形菌门、厚壁菌门和拟杆菌门组成^[38-41]。Gohar等^[42]采用高通量测序技术分析鸡油菌(C. cibarius)子实体，发现其内部细菌群落主要以变形菌门(63.2%)和拟杆菌门(20.0%)为优势种群。Kumari等^[43]通过基因组指纹法(BOX-PCR fingerprinting)分析了喜马拉雅山脉鸡油菌(Cantharellus spp.)子实体中的优势菌属，其中革兰氏阴性菌如哈夫尼菌属(Hafnia，26%)、黄单胞菌属(Stenotrophomoas，16%)和假单胞菌属(Pseudomonas，16%)占比较高。Li等^[26]通过高通量测序发现，多个地区的松茸(T. matsutake)子实体以变形菌门、拟杆菌门和厚壁菌门为优势细菌。Liu等^[44]通过高通量测序发现，微球菌目(Micrococcaleus)、芽孢杆菌目(Bacillales)、鞘氨醇单胞菌目(Sphingomonadales)和柄杆菌目(Caulobacterales)是松茸(T. matsutake)子实体不同隔室中的优势细菌菌目。Ma等^[45]通过高通量测序法分析桑黄菌(Sanghuangporus)子实体中的细菌多样性，共检测到1 476个OTUs，分为33门82纲195目355科601属679种，其中变形菌门、拟杆菌门及厚壁菌门为优势菌门，甲基娇养杆菌属(Methylotenera，8.65%)和甲基单胞菌属(Methylomonas，6.62%)为优势菌属。Oh等^[46]从240个松茸(T. matsutake)子实体切片中分离出110株细菌，并对16株代表性细菌进行16S rRNA基因鉴定，结果显示变形菌门(Proteobacteria，43.8%)、放线菌门(Actinobacteria，31.2%)和厚壁菌门(Firmicutes，25.0%)的种类占比最高；沙雷氏菌属(Serratia，34.5%)、居真菌细菌属(Mycetocola，17.3%)和假单胞菌属(Pseudomonas，16.4%)为主要菌群。王冉等^[47]采用纯培养法从云南干巴菌(T. ganbajun)子实体中分离出282株细菌，分属2门12属15种，其中假单胞菌属为主要菌群。Varese等^[48]发现厚环乳牛肝菌(Suillus grevillei)子实体中以假单胞菌属和芽孢杆菌属等为主要菌群。李云龙等^[15]从夏块菌(T. aestivum)子实体中分离筛选出233株细菌，其中假单胞菌属(40%)和布丘氏菌属(Buttiauxella，24%)为优势菌属。不同类型子实体中的细菌优势群落见表1。

2.2 子实体伴生放线菌

在子实体的生长发育过程中，也伴随着大量的伴生放线菌。放线菌G+C含量高，属于革兰氏阳性菌。在所有微生物中，放线菌，尤其是链霉菌属，是大多数活性天然产物的生产者，约2/3的已知天然抗生素被报道由链霉菌产生^[49]。许多临床使用的抗生素如头孢菌素、蒽环类、大环内酯类、糖肽类、脂多糖类和氨基糖苷类，均为放线菌产生的活性代谢产物。因此，挖掘各类子实体中放线菌的多样性对于生物资源的开发具有重要意义。Antony‐Babu等^[38]发现黑松露中放线菌门(Actinobacteria，0.6%-20.0%)为优势菌群，在外生菌根部分，链霉菌属(Streptomyces)和热嗜油菌属(Thermoleophilum)显著富集。Gohar等^[42]采用高通量测序法从鸡油菌(C. cibarius)子实体中发现优势放线菌门(Actinobacteria，6.7%)在外层的丰度明显高于内层。Varese等^[48]从厚环乳牛肝菌(S. grevillei)子实体中分离到链霉菌属(Streptomyces)为优势菌。Xiang等^[17]从双孢蘑菇(A. bisporus)子实体中分离出55株细菌和放线菌，其中放线菌(actinomycete，16.4%)为优势菌群。对16株代表性菌株进行分子鉴定，5株革兰氏阴性菌分为产碱菌属(Alcaligenes)和假单胞菌属，11株革兰氏阳性菌分为芽孢杆菌属(Bacillus)、赖氨酸芽孢杆菌属(Lysinibacillus)、类芽孢杆菌属(Paenibacillus)、潘多拉菌属(Pandorea)和链霉菌属(Streptomyces)。张兆娟等^[18]通过纯培养法对中国甘肃省、贵州省和吉林省的网柄菌子实体中的放线菌多样性进行了研究，以微杆菌属(Microbacterium)为主要菌属；从贵州省的网柄菌中分离得到17株放线菌，其中以氧化微杆菌(M. oxydans)为优势种。

2.3 子实体伴生真菌

在真菌子实体中，如桑黄、牛肝菌、肉齿菌等，除了宿主真菌外，还伴随着其他真菌生物群落的存在。Ma等^[45]利用高通量测序技术分析了桑黄(Sanghuangporus)子实体中的伴生真菌多样性，共检测到865个OTUs，分为11门34纲84目186科328属，其中优势菌门为担子菌门、子囊菌门和壶菌门，桑黄菌(Sanghuangporus)的占比最高，达到了70%。王冉等^[47]从云南干巴菌(T. ganbajun)子实体中共分离得到114株真菌，通过内转录间隔区(internal transcribed spacer, ITS)序列鉴定，结果表明所有分离真菌隶属于2门10属10种，其中子囊菌门(62%)和担子菌门(38%)为优势菌门，扁孔腔菌属(Lophiostoma)和白色星耳酵母菌属(Asterotremella)为优势菌属。李强等^[6]采用纯培养法，从西藏、吉林的新鲜翘鳞肉齿菌(S. imbricatus)子实体中共分离出26株真菌，隶属于14个属，其中球菌属(Phaeomoniella，26.9%)和隐球菌属(Cryptococcus，11.5%)为优势菌属。李鹏等^[50]同样采用纯培养法，从冬小包脚菇(Volvariella brumalis)子实体中共分离出50株真菌，通过ITS序列鉴定被划分为3门13科16属，另有2株未分类菌株，其中毛霉属(Mucor)和镰孢菌属(Fusarium)为优势菌属。从秦巴山区的牛肝菌(Boletus sp.) KL-1子实体中分离得到2株伴生真菌，经ITS序列鉴定分别为黄瘤孢菌属(Hypompces)和毛霉菌属(Mucor)的真菌^[51]。岳万松等^[52]从云南的小美牛肝菌(Boletus speciosus Frost)、铜色牛肝菌(B. aereus)、美味牛肝菌(B. edulis)和黄色疣柄牛肝菌(Leccinum crocipodium)等4种牛肝菌新鲜子实体中分离得到5株真菌，通过ITS序列鉴定分别为毛栓菌(Trametes hirsuta)、金色毛壳菌(Chaetomium aureum)、假丝酵母菌(Candida sp.)、被孢霉(Mortierella sp.)及散囊菌 (Eurotium sp.)。杨大智等^[53]从干巴菌(T. ganbajun)子实体中分离出6种伴生真菌，经ITS序列鉴定为毛霉菌属(Mucor)、赤霉菌属(Gibberella)、链格孢属(Alternaria)、拟盘多毛孢属(Pestalotiopsis)、肠茸毛菌属(Enterobryus)以及双足囊菌属(Dipodascus)真菌。

3 子实体伴生菌的生物活性

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3.1 子实体伴生菌对宿主真菌的影响

真菌子实体中存在大量伴生微生物，而真菌在发育成子实体的过程中，这些伴生菌与宿主真菌联系最为紧密，能够提高宿主真菌的抗性，同时影响其生长和代谢^[54-55]。

3.1.1 子实体伴生菌对宿主真菌生长发育的影响

子实体伴生菌能够显著影响宿主真菌的生长发育，包括菌丝生长^[56]、菌丝分支^[54]、孢子数量^[57]和生物量^[58]等。Ayudhya等^[56]从红根须腹菌[Rhizopogon roseolus (Corda) Th. M. Fr.]子实体中分离出6株变形菌门(Proteobacteria)细菌，这些细菌均能促进子实体真菌的菌丝生长。Oh等^[46]从松茸(T. matsutake)中分离出16种细菌，其中橙色迪茨氏菌(Dietzia aurantiaca)能够促进宿主真菌的生长，而乳白居真菌细菌(M. lacteus)则表现出抑制作用。Ma等^[57]从竹黄菌(Shiraia sp.)子实体中分离出的黄褐假单胞菌(P. fulva) SB1能够使竹黄菌(Shiraia sp.) S9的菌丝变得更细，并且大多数菌丝被细菌破坏成零散片段，同时，真菌的分生孢子数量显著增加；在液体培养中，竹黄菌S9与黄褐假单胞菌SB1共培养后，真菌菌球收缩，直径明显变小；扫描电镜观察发现，黄褐假单胞菌SB1与竹黄菌S9发生了紧密的物理接触，大多数细菌细胞完全附着真菌菌丝表面，部分细菌细胞几乎与真菌菌丝完全融合在。Xu等^[54]发现，从竹黄中分离的蜡样芽孢杆菌(Bacillus cereus) No.1能显著抑制竹黄菌的菌丝生长和分生孢子分化，同时增加菌丝分支数量并缩短分支距离。季红瑶^[58]从野生竹黄中分离出29株伴生真菌，并与竹黄菌(Shiraia sp.) S9进行共培养，通过平板拮抗实验筛出9株伴生真菌能够显著抑制竹黄菌的菌丝生长；此外，2株伴生真菌[亚黑管菌(Bjerkandera sp.) PTY1和炭疽菌(Colletotrichum) MAFF]在液体共培养中也能显著抑制竹黄菌的生长。

3.1.2 子实体伴生菌对宿主真菌次生代谢的影响

一些真菌在发育成子实体过程中，能够分泌活性次级代谢物来增强对宿主植物的侵染能力或抵抗外界环境，而子实体伴生菌能够显著影响宿主的相关次级代谢。Splivallo等^[41]从白松露(T. borchii)子实体中分离出α-变形菌纲和β-变形菌纲细菌，这些细菌可将非挥发性前体物质转化为挥发性化合物(如噻吩衍生物)，从而促进松露子实体香气的形成。Ma等^[57]报道了子实体伴生细菌黄褐假单胞菌(P. fulva) SB1能够刺激竹黄菌(Shiraia sp.) S9合成竹红菌素(hypocrellins)，共培养后胞内和胞外的竹红菌甲素(hypocrellin A, HA)积累显著增加，总HA产量达到225.34 mg/L，是单独培养的3.25倍。此外，竹红菌素的合成关键酶[如聚酮合酶(polyketide synthase, PKS)、锌指转录因子(zinc finger transcription factor, ZFTF)、超家族转运蛋白(major facilitator superfamily, MFS)和结合盒式转运蛋白(ATP-binding cassette transporter, ABC)]表达水平显著提高；假单胞菌SB1的活菌、热水浸提物、粗多糖提取物及透析物均能刺激HA的合成，其中活菌的诱导效果最佳；扫描电镜观察发现，假单胞菌SB1能够破坏真菌细胞膜，从而促进胞外HA的积累^[57]。Li等^[59]发现伴生黄褐假单胞菌(P. fulva) SB1通过分泌小信号分子胞外ATP (extracellular adenosine 5′-triphosphate, eATP)、钙离子(Ca²⁺)和活性氧(reactive oxygen species, ROS)促进竹黄菌(Shiraia sp.) S9竹红菌素的合成。Xu等^[54]发现从竹黄子实体中分离的蜡样芽孢杆菌(B. cereus) No. 1能显著抑制竹黄菌(Shiraia sp.) S9竹红菌素的合成，但其挥发物却能促进竹红菌素的积累，在固体培养中HA产量达到1.28 mg/cm²，是未处理组的4.06倍；进一步研究发现，蜡样芽孢杆菌No. 1能够通过还原降解竹红菌素。季红瑶^[58]从野生竹黄菌子实体中分离出29株伴生真菌，并与竹黄菌(Shiraia sp.) S9进行共培养，通过平板拮抗实验筛出19株能够促进竹红菌素合成的真菌和10株抑制其合成的真菌，通过进一步液体培养筛选出镰孢菌属真菌(Fusarium sp.) SF12对竹红菌素的促进效果最为显著，HA产量是单一培养的1.56倍，该菌通过上调竹红菌素合成关键酶基因的表达，从而提高HA产量。

3.2 抗菌及其他活性

子实体中的伴生菌本身具有广泛的抗菌活性(表2)。Xiang等^[17]从双孢蘑菇(A. bisporus)子实体中分离出55株细菌，其中Lysinibacillaceae、类芽孢杆菌科(Paenibacillaceae)、假单胞菌科(Pseudomonadaceae)和链霉菌科(Streptomycetaceae)细菌表现出广谱抑菌活性。分离出的芽孢杆菌属(Bacillus)对致病细菌和真菌均具有抑菌活性，类芽孢杆菌(B. cereus-like) DY17表现出较强的抗菌活性。Sbrana等^[60]从白松露(T. borchii)子实体中分离出259株菌株，其中101株表现出抗真菌活性，主要是放线菌(actinomycete)和假单胞菌(pseudomonad)。从松茸(T. matsutake)子实体中分离出的美洲爱文氏菌(Ewingella americana)、内生假单胞菌(P. endophytica)、沙雷氏菌、乳白居真菌细菌(M. lacteus)和奈氏西地西菌(Cedecea neteri)表现出广谱抗真菌活性，而其余11种细菌的抗菌谱较窄，其中美洲爱文氏菌几乎对所有霉菌都有抑制作用^[46]。从双孢蘑菇(A. bisporus)分离出的伴生细菌，如赖氨酸芽胞杆菌科(Lysinibacillaceae)、假单胞菌科(Pseudomonadaceae)和链霉菌科(Streptomycetaceae)等分离株对检测的病原菌表现出抗菌活性^[46]。Masrukhin等^[61]从草菇(Volvariella volvacea)中分离出3株具有抗菌活性的芽孢杆菌(Bacillus sp. C2.2、C3.8、D3.3)，通过共培养法和圆盘扩散法测试其对3种植物病原真菌[莴苣尾孢菌(Cercospora lactucae)、胶孢炭疽菌(C. gloeosporioides)和尖孢镰刀菌镰 (F. oxysporum)]的抗菌活性，其中C3.8表现出最强的抗菌活性，分析菌株C3.8的代谢产物谱发现，主要活性成分为邻苯二甲酸双酯(2-乙基己酯)(图1)。张俊兰^[62]从双孢蘑菇(A. bisporus)等6种食用菌子实体中分离出58株伴生菌，其中15株表现出抗细菌和真菌活性；抗菌活性最强的菌株为假单胞菌(Pseudomonas sp.) AB-114，该菌株通过分泌生物碱类物质来发挥抗菌作用；此外，这些菌株往往还会分泌大量的其他活性蛋白等代谢物。Oh等^[46]从松茸(T. matsutake)子实体中分离得到了16株产酶细菌，其中3株能分泌几丁质酶，7株显示出纤维素酶活性，6株具有蛋白酶活性，7株具有脂肪酶活性，此外，沙雷氏菌属和嗜麦芽寡养单胞菌(Stenotrophomonas maltophilia)具备分泌上述多种酶的能力。Bedini等^[63]从白松露(T. borchii)子实体中分离出600株细菌，其中假单胞菌属(Pseudomonas，50%)为优势菌属，检测发现90株假单胞菌包括荧光假单胞菌(P. fluorescens)、皱纹假单胞菌(P. corrugata)和托拉斯假单胞菌(P. tolaasii)能合成吲哚-3-乙酸(图1)，30株假单胞菌具有抗病原真菌的活性和合成荧光铁载体的能力，而其他菌株还能分泌蛋白酶、胶原酶和羧甲基纤维素酶。Zhou等^[64]发现从竹黄菌(Shiraia sp.) S9中分离出的伴生黄褐假单胞菌(P. fulva) SB1合成的胞外多糖EPS-1 (图1)(浓度范围为12.5-300 mg/L)对羟自由基具有清除活性(清除率为11.3%-18.1%)。桦褐孔菌[Inonotus obliquus (Fr.) Pilats]子实体中的枝顶孢属伴生菌Acremonium sp.的菌丝和发酵液乙醇提取物均展现出显著的抗肿瘤作用^[65]，伴生菌的菌丝体提取物对胃癌细胞NCI-H460和胃腺癌细胞SGC7901的抑制率分别为41.32%和36.26%，而伴生菌发酵液的抑制率则分别为56.81%和46.63%。Chen等^[66]发现解淀粉芽孢杆菌(B. amyloliquefaciens sp.) MD-b1分泌的胞外多糖能够显著抑制胃癌细胞MC-4和SGC-7901的生长。

4 总结与展望

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本文对子实体伴生菌的多样性和生物活性进行了综述，归纳了真菌子实体内微生物群落结构的主要研究方法，包括纯培养法和非培养法。通过这些方法揭示了各类真菌子实体内微生物(包括真菌、细菌和放线菌)的群落分布特征，同时探讨了各类子实体伴生菌的生物功能，特别是它们对宿主的影响以及伴生菌自身的生物活性。目前，被挖掘和报道的子实体及其伴生菌合成的活性物质仅占微生物活性成分的一小部分。由于只有少数子实体能够进行人工培育，因此未来的研究可以集中于以下几个方面：(1) 利用纯培养法分离子实体伴生菌，并挖掘能够帮助宿主真菌子实体发育、分化的伴生菌，为药用和食用菌的人工栽培提供有益帮助；(2) 基于非培养法分析子实体菌群，通过查阅文献分析菌株的亲缘关系，通过优化已报道的近缘菌株的培养条件，尝试进行非培养菌株的生长研究，以探究这部分菌群对真菌子实体发育的影响，并发现隐藏在子实体内部的功能性菌群；(3) 采用不同的提取方法获取子实体或者伴生菌的活性代谢物组分，并通过不同的药理或活性模型探究它们的生物功能，再结合分离纯化的方法，对主要的活性成分进行结构表征，结合药效筛选实验，深入挖掘子实体伴生菌的活性潜能；(4) 探讨伴生菌对子实体真菌活性成分的生物合成的调节机制，建立伴生菌-真菌共培养方法，为真菌重要活性代谢物的生物技术生产提供新型的诱导调节技术。

基金

收起

国家自然科学基金(82073955)
国家自然科学基金(81773696)
江苏省高校优势学科建设项目(PAPD)

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2025年第65卷第4期

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doi: 10.13343/j.cnki.wsxb.20250010

接收时间：2025-01-05
首发时间：2026-02-06
出版时间：2025-04-04

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收稿日期：2025-01-05
录用日期：2025-02-11

基金

National Natural Science Foundation of China(82073955)

国家自然科学基金(82073955)

National Natural Science Foundation of China(81773696)

国家自然科学基金(81773696)

Priority Academic Program Development of Jiangsu Higher Education Institutes(PAPD)

江苏省高校优势学科建设项目(PAPD)

作者信息

^1.苏州大学药学院，江苏苏州

^2.苏州大学金螳螂建筑学院，江苏苏州

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2种不同金属材料的力学参数

科 Family	属数 Number of genus	种数 Number of species	占总种数比例 Percentage of total species (%)	属 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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Fruiting body types	Dominant strain (phylum/genus)	Analytic method	References
Black truffle	Proteobacteria Bacteroidetes	High-throughput sequencing	[38]
Cantharellus spp.	Hafnia Stenotrophomoas Pseudomonas	BOX-PCR fingerprinting	[43]
C. cibarius	Proteobacteria Bacteroidetes	High-throughput sequencing	[42]
Tricholoma matsutake	Proteobacteria Bacteroidetes Firmicutes	High-throughput sequencing	[26]
T. matsutake	Micrococcales Bacillales Caulobacter	High-throughput sequencing	[44]
Sanghuangporus	Proteobacteria Bacteroidota Firmicutes	High-throughput sequencing	[45]
T. matsutake	Proteobacteria Firmicutes	Culture-dependent method	[46]
T. ganbajun	Pseudomonas	Culture-dependent method	[47]
Suillus grevillei	Pseudomonas Bacillus	Culture-dependent method	[48]
T. aestivum	Pseudomonas Buttiauxella	Culture-dependent method	[15]

Fruiting body types

Dominant strain (phylum/genus)

Analytic method

References

Black truffle

Proteobacteria

Bacteroidetes

High-throughput sequencing

[38]

Cantharellus spp.

Hafnia

Stenotrophomoas

Pseudomonas

BOX-PCR fingerprinting

[43]

C. cibarius

Proteobacteria

Bacteroidetes

High-throughput sequencing

[42]

Tricholoma matsutake

Proteobacteria

Bacteroidetes

Firmicutes

High-throughput sequencing

[26]

T. matsutake

Micrococcales

Bacillales

Caulobacter

High-throughput sequencing

[44]

Sanghuangporus

Proteobacteria

Bacteroidota

Firmicutes

High-throughput sequencing

[45]

T. matsutake

Proteobacteria

Firmicutes

Culture-dependent method

[46]

T. ganbajun

Pseudomonas

Culture-dependent method

[47]

Suillus grevillei

Pseudomonas

Bacillus

Culture-dependent method

[48]

T. aestivum

Pseudomonas

Buttiauxella

Culture-dependent method

[15]

Host fungi	Associated microbes	Activities	References
Agaricus bisporus	Bacillus	Anti-bacteria/fungi	[16]
Tuber borchii	Actinomycete Pseudomonad	Anti-fungi	[59]
Tricholoma matsutake	Ewingella americana Pseudomonas endophytica Serratia marcescens Mycetocola lacteus Cedecea neteri	Anti-fungi	[45]
Volvariella volvacea	Bacillus sp. C2.2, C3.8, D3.3	Antifungal activities against Cercospora lactucae, C. gloeosporioides, Fusarium oxysporum	[60]
A. bisporus	Pseudomonas sp. AB-114	Anti-bacteria/fungi	[61]
T. matsutake	Stenotrophomonas maltophilia	Chitinase, endoglucanase, β-glucosidase, protease, lipase activities	[45]
T. borchii	Pseudomonas	Indole-3-acetic acid, siderophores, protease, collagenase, carboxymethyl cellulose activities	[62]
Shiraia sp. S9	P. fulva SB1	Antioxidation	[63]
Inonotus obliquus	Acremonium sp.	Antitumor	[64]

Host fungi

Associated microbes

Activities

References

Agaricus bisporus

Bacillus

Anti-bacteria/fungi

[16]

Tuber borchii

Actinomycete

Pseudomonad

Anti-fungi