接上篇：https://www.chanhua.net/news.php?nid=1709

4　组学进展

　　截至目前,Genbank 数据库中已公布了蝉花3 个菌株的基因组测序结果,最早是2016 年浙江泛亚生物医药的蝉花菌株BA-001 的基因组测序结果(登录

号GCA_001644785. 1),其菌株基因组全长34. 52Mb,基因组覆盖度为28 × ,GC 含量52. 5% [68] 。2018年中国科学院植物生理生态研究所王成树团队提交蝉花菌株CC02 基因组数据( 登录号GCA _002968875. 1),其基因组全长为34. 11 Mb,基因组覆盖度为80 × ,GC 含量53. 0% ,含有9 701 个蛋白编码基因[69] 。2020 年安徽农业大学对蝉花菌株ZJ1611 进行了基因组测序( 登录号: GCA _010211705. 1),其全长为33. 8 Mb,基因组覆盖度为233 × ,GC 含量为53. 1% ,含有11 106 个蛋白编码基因[70] 。LU 等[69] 通过基因组和比较基因组分析发现蝉花编码的蛋白家族属于典型的虫生真菌蛋白,包括侵染寄主的蛋白酶和几丁质酶等,并发现蝉花无性子实体的产生与交配型(mating-type,MAT)位点控制无关。线粒体基因组作为研究物种起源和进化的有力工具,FAN 等[71] 测定蝉花3 个菌株CCAD02、ARSEF11726和BA-001 的线粒体基因组,其线粒体基因组总长分别为56 581、49 138 和约56 080 bp, 都含有2 个核糖体RNA 和25 个转运RNA,内含子数量分别为25、19 和25,内含子数量的差异是导致线粒体基因组长度差异的原因;通过种间分析确定蝉花的分类位置,从系统发育树上揭示了蝉花与蛹虫草亲缘关系相近。

图1　蝉花生物活性作用

Fig. 1　The biological activities in Chanhua

注:图中图片来源于网络。

　　代谢组学采用气相色谱质谱联用、高效液相色谱质谱联用、核磁共振等技术及数据分析处理方法,分析由于生物或非生物因素导致的样品中小分子代谢物质成分变化。近年来,代谢组学在蝉花中的应用主要集中在分析不同部分、不同培养时间或不同产地的代谢成分变化。如HE 等[38] 对野生蝉花、人工柞蚕蛹培养和人工固体培养基的蝉花菌丝体的代谢组学分析发现在野生虫草和人工柞蚕蛹培养蝉花的虫体部分存在着高水平的非核糖体多肽( nonribosomal peptides,NRPs),鞘脂类的水平在野生蝉花和人工柞蚕蛹培养的蝉花无论是在虫体部分还是在孢梗束部分都高于蝉花菌丝体,除了NRPs 和鞘脂外,虫体部分含量较高的还有红土霉素、茴香霉素、链霉胺酮和乌司他洛辛。孢梗束和菌丝体部分含量较高的为生育酚、3′-脱氧花青素、γ-氨基丁酸和磷脂等有益的物质[38] 。张万存等[36] 对不同培养时间下蝉花菌丝体代谢组分析发现,随着培养时间不同代谢物具有显著变化,培养前期产生较多的细胞松弛素E、肌醇、γ-氨基丁酸、乙酰肉碱和亚油酸等与侵染宿主、早期信号传导和氧化供能有关的物质;中期产生较多的甘露醇、海藻糖、亚麻酸、柠檬酸、丙酮酸和7,8-二羟基油酸等与抗逆和氧化供能有关的物质,此外红酵母红素、δ-氨基-γ-酮戊酸2 种抗生素含量也达到了最大值;培养后期产生较多的脯氨酸、2,4-二氨基丁酸、3′-deoxy-hanasanagin、ulvaline、赤霉素和烟酰胺等与抗逆、抗氧化、产孢和细胞保护有关的物质。何亚琼等[37] 对人工培养柞蚕蝉花的不同部分代谢组学分析表明,不同部位的代谢物显著不同,蝉花虫体体内物质以大分子储藏物质为主,虫体表面菌丝中有大量次生代谢产物,孢梗束的次生代谢产物相对较少。

　　转录组学技术是寻找功能基因,揭示天然活性化合物生物合成及调控机理的重要工具。目前利用转录组学技术在阐明蝉花的活性物质如虫草素、麦角甾醇等的生物合成、揭示活性物质的作用机理等方面均有报道。LIU 等[72] 通过转录组学技术挖掘蝉花中与虫草素合成相关的候选基因,并发现5′-核苷酸酶和腺苷脱氨酶参与虫草素的合成。SU 等[73] 通过转录组揭示了蝉花中作为氮源的尿素增加麦角甾醇合成的调节机理。朱伟坚等[60] 通过转录组揭示了蝉花中HEA 是通过激活腺苷A1 R,下调A2 AR 及调控一系列疼痛相关基因发挥其镇痛作用。

　　蛋白质组学作为后基因时代的标志之一,是药物发现及开发、寻找靶标蛋白等的有效途径。目前关于蝉花的蛋白质组学的文献报道较少。WANG 等[55] 通过蛋白质组学和飞行时间质谱分析研究蝉花对肝细胞抗癌机制,发现蝉花水体物通过G2 / M 细胞周期阻滞抑制人肝癌细胞系MHCC97H 的生长。

5　前景与展望

　　蝉花、冬虫夏草、蛹虫草是我国虫草类真菌的主要代表物种,其中蝉花的活性成分与我国名贵虫草———冬虫夏草相似,有望作为冬虫夏草的替代品,市场前景广阔,且蝉花部分产品已经实现了商品化。2021 年,随着蝉花人工栽培子实体批准为新食品原料,使得蝉花的应用范围进一步扩大,且随着蝉花中活性物质成分多糖、HEA、麦角甾醇、多球壳菌素等功能活性的探明、分离提取纯化工艺等的完善,蝉花的应用价值及商业化应用将会进一步扩大,市场需求也将会急剧增加。面对这一现状,在已有的研究基础上,未来应加强蝉花的人工培养技术研究,形成产业化、规模化蝉花的替代寄主昆虫人工培养方式,进一步加强固体培养技术和液体发酵培养技术;加快蝉花质量标准的制定,如活性物质的检测方法、含量的标准;加快蝉花专利的申请,为蝉花的规范化、产业化、市场化应用提供支撑;对具有生理活性的物质成分应加强分离纯化及生物安全性评估,为其应用开发奠定基础;同时应加强、加深蝉花生物活性机理的研究,蝉拟青霉菌株资源的收集、评价、鉴定及采用分子生物学技术等手段对菌株进行改造等基础研究工作,为蝉花在食品、药品、化妆品等多领域的开发应用提供研究基础和科学依据。

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Research progress and analysis of Cordyceps chanhua

ZHOU Sijing1,2,3 , QIAO Yuchen1 , LIU Guijun1∗ , CHEN Wei2,3 ,WANG Ping1 , SONG Meifang1 ,WANG Chengtao2,3∗

　　1(Institute of Radiation Technology, Beijing Academy of Science and Technology, Beijing 100875, China)

　　2(Beijing Advanced Innovation Center for Food Nutrition and Human Health, Beijing Technology and Business University,Beijing 100048, China)3(Beijing Engineering and Technology Research Center of Food Additives, Beijing Technology and Business University, Beijing 100048, China)

　　ABSTRACT　Chanhua (Cordyceps chanhua), one of the famous Cordyceps fungi, has a long history of edible and medicinal use in China.It exhibits a high nutritional value and various medicinal properties, such as immunomodulation-related activity, anti-tumor effect, improving renal function, and so on. In this review, the progress of the chemical composition, artificial cultivation, biological functional activities,and the current omics in Cordyceps chanhua were summarized, as well as the future prospects of C. chanhua were briefly predicted,aiming to provide a reference for the future application of C. chanhua in food, medicine, health products, cosmetics, and other

fields.

　　Key words　Cordyceps chanhua; composition; biological activity; artificial cultivation; omics progress