摘要
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。
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
1 子实体伴生菌多样性分析
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
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
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
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
2 子实体伴生菌的分类
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) 1
| Fruiting body types | Dominant strain (phylum/genus) | Analytic method | References |
|---|---|---|---|
| Black truffle |
Proteobacteria Bacteroidetes | High-throughput sequencing |
[ |
| Cantharellus spp. |
Hafnia Stenotrophomoas Pseudomonas | BOX-PCR fingerprinting |
[ |
| C. cibarius |
Proteobacteria Bacteroidetes | High-throughput sequencing |
[ |
| Tricholoma matsutake |
Proteobacteria Bacteroidetes Firmicutes | High-throughput sequencing |
[ |
| T. matsutake |
Micrococcales Bacillales Caulobacter | High-throughput sequencing |
[ |
| Sanghuangporus |
Proteobacteria Bacteroidota Firmicutes | High-throughput sequencing |
[ |
| T. matsutake |
Proteobacteria Firmicutes | Culture-dependent method |
[ |
| T. ganbajun | Pseudomonas | Culture-dependent method |
[ |
| Suillus grevillei |
Pseudomonas Bacillus | Culture-dependent method |
[ |
| T. aestivum |
Pseudomonas Buttiauxella | Culture-dependent method |
[ |
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
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
3 子实体伴生菌的生物活性
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
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
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
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(
| Host fungi | Associated microbes | Activities | References |
|---|---|---|---|
| Agaricus bisporus | Bacillus | Anti-bacteria/fungi |
[ |
| Tuber borchii |
Actinomycete Pseudomonad | Anti-fungi |
[ |
| Tricholoma matsutake |
Ewingella americana Pseudomonas endophytica Serratia marcescens Mycetocola lacteus Cedecea neteri | Anti-fungi |
[ |
| Volvariella volvacea |
Bacillus sp. C2.2, C3.8, D3.3 |
Antifungal activities against Cercospora lactucae, C. gloeosporioides, Fusarium oxysporum |
[ |
| A. bisporus | Pseudomonas sp. AB-114 | Anti-bacteria/fungi |
[ |
| T. matsutake | Stenotrophomonas maltophilia | Chitinase, endoglucanase, β-glucosidase, protease, lipase activities |
[ |
| T. borchii | Pseudomonas | Indole-3-acetic acid, siderophores, protease, collagenase, carboxymethyl cellulose activities |
[ |
| Shiraia sp. S9 | P. fulva SB1 | Antioxidation |
[ |
| Inonotus obliquus | Acremonium sp. | Antitumor |
[ |
图1 一些子实体伴生菌的活性代谢物结构式
Figure 1 Structures of some bioactive metabolites from fruiting body-associated microbes.
4 总结与展望
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。
作者贡献声明
沈文浩：综述初稿撰写及修改；郑丽屏：图表制作及综述修改；周建芹：文献分析及综述修改；王剑文：提供研究思路与论文写作指导。
利益冲突
作者声明不存在任何可能会影响本文所报告工作的已知经济利益或个人关系。
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