摘要
核盘菌（Sclerotinia sclerotiorum (Lib.) de Bary）是一种具备广泛寄主，能造成至少75个科，278个属的400多种植物病害的植物病原真菌，通常能造成许多重要农作物如：向日葵、大豆、油菜、食用豆类、鹰嘴豆、干豌豆、扁豆及部分单子叶植物如：洋葱和郁金香等的严重病害。在分类学上，核盘菌属于同宗配合子囊真菌门，柔膜菌目，核盘菌科。在核盘菌的生长发育过程中，通过营养菌丝的聚集形成大量表面黑色、质地坚硬的多细胞组织结构，称之为菌核，这些结构使核盘菌在农田土壤中的存活及在传染性繁殖的持久性过程中起到非常重要的作用。菌核能在恶劣的生物及非生物环境如：低温、干燥及微生物刺激等环境中存活数年之久，在适合的环境条件下，菌核能从新萌发形成菌丝或者子囊盘，当子囊盘发育成熟时能释放出大量的子囊孢子开始新的侵染循环。由于核盘菌菌核能适应恶劣环境长期存活而保持长久致病性，所以核盘菌病害一直是世界上防治困难且每年造成巨大经济损失的农业病害之一。
菌核的生长发育及萌发过程涉及到几个不同的阶段，且受到许多来自外源环境如：光照、温度、活性氧浓度、环境pH、机械因素和营养成分等的调控，同时也受到来自内源性遗传因子的严密调控，如：一些与菌核发育相关且独立和偶联的转录调控途径，特别是一些与AMP环化酶相关的磷酸化，类似ERK的磷酸化-丝裂原活化蛋白激酶链及丝氨酸/苏氨酸磷酸酶2A/2B等的调控途径相关，而且在核盘菌生长发育过程中，菌核还扮演着从无性发育到有性发育转变的关键角色。
本研究从核盘菌的基因组和突变体库中分离得到两个与核盘菌生长发育调控相关的转录因子Ss-Nsd1和Ss-Fox1，阐明了编码GATA类IVb型锌指蛋白转录因子Ss-Nsd1和Forkhead蛋白家族类转录因子Ss-Fox1在核盘菌生长发育过程中的调控作用。转录因子Ss-Nsd1是一个与构巢曲霉的NsdD同源的蛋白，在核盘菌的子囊梗及子囊盘形成早期，Ss-nsd1是一个依赖于UV-A的组织特异性表达的基因。在Ss-nsd1的敲除突变菌株（ΔSs-nsd1）的培养过程中，其营养菌丝和菌核内部均能产生大量的分生孢子梗及分生孢子，而相反，在野生型菌株中只在菌核表面产生少量分生孢子。由于Ss-nsd1基因的敲除，影响了菌核结构发育的完整性及萌发过程中产囊体的形成，从而导致不能萌发形成子囊盘。同时研究还发现，Ss-nsd1基因的敲除菌株还影响侵染垫的形成，而Ss-fox1基因是一个编码Forkhead蛋白家族的基因。因此，在本研究中构建了Ss-nsd1和Ss-fox1基因的敲除载体，通过基因敲除研究表明Ss-nsd1基因的功能及特点与子囊菌中维持有性及无性发育平衡的功能基因nsdD相类似，主要是通过负调控分生孢子及正调控菌核的发育、侵染垫的形成及子囊盘的发育来维持核盘菌无性到有性发育的转变和平衡，而Ss-fox1的敲除菌株（ΔSs-fox1）则影响核盘菌子囊盘的发育。
未来的研究工作主要关注与Ss-nsd1和Ss-fox1基因相偶联的一些信号通路及其下游基因的相关研究，旨在搞清楚丝状真菌在有性及无性发育过程中的一些共有的和独特的信号转导途径。
Sclerotinia sclerotiorum (Lib.) de Bary has host range comprises at least75families and278genera including more than400plant species, it is a devastatingnecrotrophic fungal plant pathogen of many agriculture crops including sunflower,soybean, oilseed rape, edible dry bean, chickpea, dry pea, lentils and monocotyledonsuch as onion and tulip. Taxonomically this fungus is a homothallic ascomycetebelonging to the family Sclerotiniaceae of the order Helotiales. Developmentally, ahallmark of this and related fungi is the production of hardy multicellular sclerotiaformed from the aggregation of vegetative hyphae and enclosed by a melanized rindlayer. These structures play a significant role in the survival and persistence ofinfectious propagules in agriculture fields. Sclerotia can survive under harshbiological and physical environments including low temperature, microbially activesoils, and dry environments for several years. Under suitable environmentalconditions, sclerotia germinate either into vegetative hyphae (myceliogenicgermination) or into apothecia (carpogenic germination), with the latter releasinglarge quantities of ascospores that initiate new disease cycles. With its adaptationsfor long-term survival and pathogenicity on a broad range of hosts, S. sclerotiorum isone of the most challenging agricultural pathogens to manage and causes largeglobal economic losses annually.
The development and carpogenic germination of sclerotia involves severaldistinct stages and is affected by numerous exogenous factors such as photoperiod,temperature, oxygen concentration, ambient pH, mechanical factors and nutrients. Atthe same time, this process is also tightly regulated by intrinsic genetic factors.Independent and cross-talking pathways have been shown to be involved in sclerotium development; in particular, phosphorylative relay involving AMP cyclase,ERK-like mitogen-activated protein kinase and Ser/Thr phosphatases type2A and2B, and represention a transitional structure spanning asexual and sexualdevelopment during the S. sclerotiorum development. In this study, we isolated twotranscriptional factors Ss-Nsd1and Ss-Fox1from the S. sclerotiorum, and we reporton the regulatory functions of the S. sclerotiorum GATA-type IVb zinc-fingertranscription factor Ss-Nsd1and Ss-Fox1in these processes. Ss-Nsd1is orthologousto the Aspergillus nidulans NsdD (never in sexual development) proteins.TheSs-nsd1gene showed a strong UV-A light-dependent and tissue-specific transcriptaccumulation in stipe and early developing apothecium tissues. The Ss-nsd1deletionmutant strains (ΔSs-nsd1) produced phialides and phialospores promiscuously inboth vegetative culture and within sclerotia. In striking contrast, phialosporedevelopment occurred only on the sclerotium surface in the wild type. Loss ofSs-nsd1function affected sclerotium structural integrity and disrupted ascogoniaformation during conditioning for carpogenic germination. As a consequence,apothecium development was abolished. The Ss-nsd1deletion mutants were alsodefective in compound appressorium formation. However, the Ss-fox1is a Forkheadprotein family coding gene. In sum, We functionally and characterized the Ss-nsd1and Ss-fox1genes by gene deletion and demonstrate that the Ss-nsd1functionssimilarly as its ascomycete orthologs nsdD gene in balancing sexual and asexualdevelopments by both negatively regulating phialospore (spermatia) formation andpositively regulating sclerotium, compound appressorium and apotheciumdevelopment， however the Ss-fox1deletion mutant strains (ΔSs-fox1) affectapothecia development in S. sclerotiorum.
Further study is for Ss-nsd1and Ss-fox1genes linking signaling pathways anddownstream genes have the potential to increase understanding of the common andunique pathways adopted for varying sexual and asexual development in filamentousfungi.
引文
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