食用菌有性发育相关基因的研究进展
2016-03-24常黎明张亚军
张 育,常黎明,张亚军,谢 放
(兰州交通大学化学与生物工程学院,甘肃 兰州 730030)
食用菌有性发育相关基因的研究进展
张 育,常黎明,张亚军,谢 放
(兰州交通大学化学与生物工程学院,甘肃 兰州 730030)
摘 要:从构巢曲霉veA基因的发现出发,以研究时间先后为顺序,总结了近30 a来发现的与食用菌有性发育相关的基因及其作用,其中重点介绍了双孢菇、香菇和冬虫夏草中有性发育相关基因的功能,并对今后食用菌有性发育的研究进行了展望。
关键词:食用菌;有性发育;功能基因;综述
食用菌发育包括无性和有性两个阶段,一般以产生亲和性交配型双核菌丝为界线[1]。通常,子实体的发生与发育会受到外界环境的影响,但归根结底还是基因选择性表达的结果。早在100 a以前,人们就开始对食用菌的生物学特性、驯化育种、培养基质、栽培技术、营养生理和病虫防治等展开了研究[2],但关于食用菌有性发育分子机理方面的研究成果相对较少。2010年鲍大鹏等[3]完成了草菇的全基因组测序,该成果推动了一些珍稀食用菌如冬虫夏草、羊肚菌等的分子机理研究。随着分子生物学技术的发展,国内外学者对食用菌发育分子机理的研究也越来越深入,取得了阶段性的成果。笔者综述了近30 a来与食用菌有性发育相关的功能基因的研究进展,以期为某些不能人工栽培的珍稀食用菌的开发利用提供参考。
1 构巢曲霉中与有性发育相关的基因及作用
早在1965年,Kafer等[4]从模式真菌构巢曲霉的突变株中分离到veA基因的突变基因(veA1基因),因该基因和其他已知功能的蛋白质之间无同源性,导致该基因的研究一直停滞。1981年,Champe等[5]发现,veA蛋白在真菌中作为有性发育的正调节因子,调节着有性发育的整个过程。直至2002年,Kim等[6]发现veA突变体在适宜于野生型产生子囊果的条件下也不会产生子囊果,充分证明了veA基因的表达与有性发育的启动是直接相关的,它是有性发育的启动基因之一。这一结果的获得对食用菌有性发育机理的研究起着里程碑式的作用。
2 食用菌中与有性发育相关的基因及作用
2.1 双孢菇
食用菌中,与有性发育相关的基因最早是在双孢菇(A.bisporus)中发现的,在研究可利用培养基基质的食用菌基因时偶然发现了一些基质降解酶基因,如葡萄糖-6-磷酸脱氢酶基因、甘露醇脱氢酶基因、几丁质酶基因及鸟苷酸转移酶基因等(表1),在子实体形成过程中可差异性表达,由此推测这些酶对应的基因可能与有性发育相关。
表1中的8个基因,目前只有甘露醇脱氢酶基因,鸟苷酸转移酶基因,PPO/HSP基因的功能得到了验证,它们在有性发育过程中分别起到渗透压调节,产物分布以及调剂有性发育的作用,但其他与有性发育相关的基因的功能尚未得到验证,还需进一步研究。
2.2 香 菇
香菇中有性发育相关基因的研究最多,进展也较快。目前,已发现了11个与有性发育相关的基因,如表2所示。通过研究这些基因在子实体形成过程中表达量的变化,推测这些基因可能与有性发育相关。此外,还发现了105个未命名的基因,也可能与有性发育相关,这些基因涉及基础代谢、细胞结构、信号传导和胁迫响应等生理活动,说明子实体形成过程与多个代谢途径和信号转导相关。
表1 双孢菇(A.bisporus)中与有性发育相关的基因
表2 香菇中与有性发育相关的基因
2.3 冬虫夏草
2009年,Yao等[26]研究了子座形成初期中国被毛孢基因转型突变和蝙蝠蛾拟青霉基因的表达,发现GC和AT基因转型酶可能来自于中国被毛孢,该酶可能在促进冬虫夏草无性发育到有性发育的连接过程中起关键性作用。
2010年,Zhang等[27]克隆得到了冬虫夏草僵虫和子座两部分的cDNA文库,并将这两个文库的序列进行了聚类分析,发现在发育过程中,核酸水平存在高度的复杂性,而蛋白质水平依旧很保守;Zhu等[28]在研究冬虫夏草由无性发育向有性发育过渡时GC-AT基因转型时发现,冬虫夏草的成熟不仅与蝙蝠蛾拟青霉和冬虫夏草突变体基因子实体的增加聚集有关,而且与含有GC-AT基因转型的冬虫夏草菌基因的特异性表达有关,这个发现对明确冬虫夏草由无性型向子座形成过程起到至关重要的作用。
2011年,Zhu等[29]对有性发育过程中基因的转换和颠换及其随子实体成熟发生的变化做了进一步研究,发现基因转型可能是冬虫夏草菌子座萌发和成熟所必须的。
2012年,Gao等[30]对冬虫夏草菌基因转型在子座成熟阶段的变化进行了研究,发现冬虫夏草中至少有5个转换和颠换突变基因型同时出现,该基因型在子实体产生过程中显著性表达可能为冬虫夏草子座萌发和成熟提供重要的信息,使冬虫夏草由无性型转换为有性型;Yao等[31]研究了两种冬虫夏草独特成熟模式下GC偏差基因型的变化,结果表明冬虫夏草中含有两个特殊的GC偏差基因型,这2个GC偏差基因和冬虫夏草的其他基因型是否为内部或外部的物种突变还有待进一步的研究。
2013年,李建宏[32]构建了中国被毛孢有性发育起始阶段cDNA文库,该文库中存在一段与其他真菌有性发育相关基因有极大相似性的veA基因。
近年来,在冬虫夏草有性发育过程中的研究取得了阶段性的进展,但总体来讲,这些研究缺乏系统性,希望在以后的研究中多做一些与实际相关的研究。
2.4 其 他
有关裂褶菌发育分子机理的研究起步较早,Mulder等[33]在研究裂褶菌子实体形成过程中单核菌丝和双核菌丝中RNA的差异性表达时发现,在子实体形成初期,有性发育相关基因的表达量远低于其成熟期的表达量。Schuren等[34]研究了子实体形成过程中Sc7/Sc14基因家族编码胞外蛋白的特异性表达,发现Sc7蛋白在培养基和子实体内均存在,并可看到蛋白松散地锚定在菌丝壁表面。此外,还有一些其他食用菌中的研究成果,详见表3。
表3 其他食用菌中与有性发育机理相关的研究概况
3 展 望
20世纪80年代至今,对食用菌的研究一直没有中断过,虽然已经取得了一些阶段性成果,但还存在很多问题未能确定:(1)与有性发育相关的基因已发现较多,但其具体作用位置还有待进一步验证;(2)子实体形成是一个复杂的过程,其分化受多个基因调控,各基因之间的相互关系如何,具体怎么调控有待进一步深入研究;(3)食用菌有性发育的研究对象较分散、研究内容较笼统、研究成果不系统。总而言之,食用菌有性发育是一个系统而庞大的课题,需要各方面协同努力才能攻克这一难题。
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(责任编辑:成 平)
Research Progress on Sexual Development Related Genes in Edible Fungi
ZHANG Yu,CHANG Li-ming,ZHANG Ya-jun,XIE Fang(School of Chemistry and Biology Engineering, Lanzhou Jiaotong University, Lanzhou 730030, PRC)
Abstract:This paper reviews the sexual development related genes and their functions in edible fungi in research time sequence in recent 30 years since the discovery of Aspergillus nidulans veA gene, mainly on Agaricus bisporus, Lentinula edodes and Cordyceps sinensis, and prospects the further reseach on the edible fungi’s sexual development in the future.
Key words:edible fungi; sexual development; functional gene; review
作者简介:张 育(1989-),女,甘肃天水市人,硕士研究生,主要从事资源与环境微生物研究。
基金项目:甘肃省科技支撑基金资助项目(212256)
收稿日期:2015-10-20
DOI:10.16498/j.cnki.hnnykx.2016.01.032
中图分类号:S646.032
文献标识码:A
文章编号:1006-060X(2016)01-0115-04