接上篇：https://www.chanhua.net/news.php?nid=1695

4 蝉花的分子生物学研究进展

　　随着现代分子生物学的发展，基因功能分析已经在多种真菌上应用。利用分子生物学技术研究基因功能和相应的代谢途径，在蛹虫草真菌的研究中已经应用，但在蝉花菌中研究还很少[60]。据报道，蝉花菌的基因组大小33.9 Mb, 包含9701 个蛋白编码基因[61]。本团队采用三代测序技术，对我们从江苏宜兴采集到的蝉花菌株2-2 进行了基因组测序分析，测得的基因组大小超过34.8 M。经过组装，获得了19 个Contig，其中超过3M 的有6 个，达到染色体水平。通过分析，我们发现2-2 菌株基因组编码9942 个蛋白。

　　通过表达谱和代谢组学分析，发现不同培养阶段，不同方法，蝉花真菌细胞中的基因表达量不同，细胞中成分含量也有差异。人工培养条件下，无性态阶段与交配型基因的表达无关，说明无性态和有性态在基因表达层面是相互独立的。He 等[62]对蝉花代谢组学研究显示，与野生蝉花相比，人工培养的代谢更活跃，它们含有与三羧酸循环相关的代谢物，如肉碱、乙酰肉碱、烟酸、乙醛酸、柠檬酸、γ- 氨基丁酸和琥珀酸，而相比之下，野生的新陈代谢要弱。采用质谱等多种分析方法对破壁的分生孢子成分分析发现，分生孢子中至少含有59种挥发性成分，这些成分具有潜在的医疗保健作用[26]。

　　真菌几丁质酶在虫生真菌寄生昆虫过程中的几丁质降解、细胞壁构建、几丁质循环、营养获取、自溶和毒力等方面发挥着重要作用。Peng 等[63]结合蝉花的基因组和基因表达分析，发现蝉花基因组中有18 个糖苷水解酶基因，其中17 个是几丁质酶基因。这些基因可以分为A、B、C 等3 个组，其中A 组（包括7 个基因）的表达量大，并在蝉花生长的各个阶段都表达；B 组（包括7 个基因）在含有几丁质的基质上高表达，可能与寄生昆虫相关。

　　虫草素能够诱导肺癌细胞的凋亡，在临床上具有良好的应用前景。但其化学合成的成本高但产量低而难以实际应用。生物合成就成为非常重要的途径。Liu 等[64]利用高效液相色谱测定了虫草素在蝉花不同组织中的含量，发现在人工栽培条件下，虫草素在蝉花菌核中含量最高，在子实体中次之，在菌丝中没有检测到。蛹虫草中虫草素的生物合成是由脱氧腺苷（5 ’-deoxyadenosine）途径进行的，而蝉花菌的基因组中却没有其同源基因。Liu 等[64]利用转录组技术，对蝉花菌丝体、子实体和菌核的转录组进行了分析，并鉴定差异表达基因。在菌丝和菌核表达文库中，分别鉴定出1576 个上调基因和2300 个下调基因。在菌丝体-子实体文库和子实体-菌核体文库中，分别有1604 和1474 个基因上调，1365 和1320 个基因下调。Liu 等[64]对代谢途径分析认为，腺苷是嘌呤途径的重要中间产物，蝉花中虫草素的合成可以从腺苷开始，经过一系列反应生物合成虫草素。基因差异表达分析发现，与嘌呤途径相关的基因有19 个在菌丝与菌核中差异表达，28 个在菌丝与子实体中差异表达，16 个基因在菌丝与菌核中的差异表达。通过定量PCR 反应验证，发现有6 种关键酶基因表达差异明显。其中，参与虫草素生物合成的嘌呤核苷酸代谢途径中的5 ’-核苷酸酶和腺苷脱氨酶在菌核中显著上调，与虫草素在组织中的含量动态相符，推测虫草素是以嘌呤途径中的重要中间产物次黄嘌呤单核苷酸（IMP）为底物生物合成虫草素的关键酶[65]。

5 问题与展望

　　近年来随着人们对健康生活需求的提高，食药用真菌的研究开发越来越受到重视。作为我国重要的食药用菌，蝉花的研究也取得了很多成果，研究不断深入，但与当前人们的需求还有很大差距。虽然目前认为蝉花与冬虫夏草的许多功能相似，但蝉花与冬虫夏草、蛹虫草研究的科学性、规范性相比，还明显不足[2]；对蝉花的有效药用成分和功效了解不足，还没有科学的标准对蝉花品质进行科学评价，目前采用虫草素、虫草酸等成分对冬虫夏草、蛹虫草的评价体系不适合蝉花，由此造成对人工栽培蝉花的质量标准评价不科学；对蝉花的生物学、生态学和与寄主互作关系认识不足，缺乏优质的栽培技术，如有研究认为栽培基质中添加4%的蝉蛹可以提高蝉花的代谢产物[13]，而有研究认为蝉花生长环境中细菌群体结构影响蝉花代谢[66-67]，不同发酵条件影响代谢产物的种类和含量等，制约了其在各个领域的进一步开发和应用。相对于冬虫夏草、蛹虫草等，蝉花产业相对较小。天然蝉花资源十分有限，且由于对野生资源认识不足，经常将其他菌体误认为蝉花，或因为野外卫生条件不良，而造成负面影响。人工培养将是重要发展方向。蝉花的新功能有待进一步开发，如功能食品、化妆品等方面等。新的加工技术的应用将使产品形式多样化[68]。虽然人工培养的蝉花菌体已经是新资源食品，但是执法实践需要在产业发展中进一步完善。

　　根据目前蝉花的生产和研究现状，必须用现代前沿生物科学技术，促进蝉花的科学研究，为蝉花可持续高质量的科学应用提供保障。随着蝉花分子转化系统的建立[63]，要将功能基因组学、转录组学、蛋白组学、代谢组学、脂组学等前沿科学技术应用于蝉花的研究，明确蝉花的分子遗传特性和基因调控网络，深入揭示蝉花的生物学特性，明确活性成分的代谢网络；将蝉花的生物学研究与医疗保健效果研究紧密结合，明确蝉花组分的医疗保健作用，为蝉花的科学应用提供依据；在此基础上，发展基因编辑技术，消除蝉花基因组中可能的不良产物基因，提高有用的活性成分和营养组分，为满足人们对蝉花日益增长的需求提供技术支持；将蝉花与其他食药用菌进行（近）远缘杂交[69]，结合分子育种，开发优质新品种。随着研究的快速深入发展，蝉花必将在新药研发和人们健康生活中起到重要作用。

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