Article(id=1226554096795172996, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226554095926952065, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20250006, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1735747200000, receivedDateStr=2025-01-02, revisedDate=null, revisedDateStr=null, acceptedDate=1740326400000, acceptedDateStr=2025-02-24, onlineDate=1770362884948, onlineDateStr=2026-02-06, pubDate=1751558400000, pubDateStr=2025-07-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1770362884948, onlineIssueDateStr=2026-02-06, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1770362884948, creator=13701087609, updateTime=1770362884948, updator=13701087609, issue=Issue{id=1226554095926952065, tenantId=1146029695717560320, journalId=1192105938417971205, year='2025', volume='65', issue='7', pageStart='2771', pageEnd='3233', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1770362884741, creator=13701087609, updateTime=1770363575040, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1226556991309529548, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226554095926952065, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1226556991309529549, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226554095926952065, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=2854, endPage=2863, ext={EN=ArticleExt(id=1226554097101357193, articleId=1226554096795172996, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Research progress in specific subunits of RNA polymerase III in Saccharomyces cerevisiae, columnId=1192149543727808575, journalTitle=Acta Microbiologica Sinica, columnName=Review, runingTitle=null, highlight=null, articleAbstract=

Saccharomyces cerevisiae is a classic model organism for studying the biochemical mechanisms of eukaryotic cells. Eukaryotes have three main types of RNA polymerases: RNA polymerase I (RNAPI), RNA polymerase II (RNAPII), and RNA polymerase III (RNAPIII). Among them, RNAPIII has the most complex structure, consisting of 17 subunits, and it is primarily responsible for the synthesis of transfer RNA (tRNA). Compared with RNAPII consisting of 12 subunits, RNAPIII contains a unique heterotrimer Rpc82/31/34 and a heterodimer Rpc53/37 which is homologous to the counterpart of RNAPI. This paper reviews the structures and functions of the specific heterotrimer and heterodimer in RNAPIII, aiming to lay a theoretical foundation for further studies on the modification mechanisms and assembly processes of specific subunits of RNAPIII in S. cerevisiae.

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酿酒酵母(Saccharomyces cerevisiae)是研究真核生物细胞生化机制的经典模式生物。真核生物中主要存在3种RNA聚合酶(RNA polymerase),分别为RNA聚合酶Ⅰ (RNAPI)、RNA聚合酶Ⅱ (RNAPII)和RNA聚合酶Ⅲ (RNAPIII)。其中,RNA聚合酶Ⅲ的结构最为复杂,包含17个亚基,主要负责合成转运RNA (transfer RNA, tRNA)。相较于包含12个亚基的RNAPII,RNAPⅢ含有一组独特的异三聚体Rpc82/31/34,以及一对与RNAPI同源的异二聚体Rpc53/37。本文综述了RNAPIII中异三聚体和异二聚体的结构与功能,为深入研究酿酒酵母RNAPⅢ特异亚基修饰机制和组装过程提供了理论依据。

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作者贡献声明

杨欣怡:文献的检索与归纳,撰写全文及修改;李盼:全文指导及修改;曾凡力:获取基金,全文指导及修改。

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酿酒酵母RNA聚合酶特异亚基的研究进展
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杨欣怡 , 李盼 , 曾凡力
微生物学报 | 综述 2025,65(7): 2854-2863
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微生物学报 | 综述 2025, 65(7): 2854-2863
酿酒酵母RNA聚合酶特异亚基的研究进展
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杨欣怡, 李盼, 曾凡力
作者信息
  • 河北农业大学 生命科学学院,河北 保定
Research progress in specific subunits of RNA polymerase III in Saccharomyces cerevisiae
Xinyi YANG, Pan LI, Fanli ZENG
Affiliations
  • College of Life Sciences, Hebei Agricultural University, Baoding, Hebei, China
出版时间: 2025-07-04 doi: 10.13343/j.cnki.wsxb.20250006
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酿酒酵母(Saccharomyces cerevisiae)是研究真核生物细胞生化机制的经典模式生物。真核生物中主要存在3种RNA聚合酶(RNA polymerase),分别为RNA聚合酶Ⅰ (RNAPI)、RNA聚合酶Ⅱ (RNAPII)和RNA聚合酶Ⅲ (RNAPIII)。其中,RNA聚合酶Ⅲ的结构最为复杂,包含17个亚基,主要负责合成转运RNA (transfer RNA, tRNA)。相较于包含12个亚基的RNAPII,RNAPⅢ含有一组独特的异三聚体Rpc82/31/34,以及一对与RNAPI同源的异二聚体Rpc53/37。本文综述了RNAPIII中异三聚体和异二聚体的结构与功能,为深入研究酿酒酵母RNAPⅢ特异亚基修饰机制和组装过程提供了理论依据。

酿酒酵母  /  RNA聚合酶III  /  特异亚基  /  转录

Saccharomyces cerevisiae is a classic model organism for studying the biochemical mechanisms of eukaryotic cells. Eukaryotes have three main types of RNA polymerases: RNA polymerase I (RNAPI), RNA polymerase II (RNAPII), and RNA polymerase III (RNAPIII). Among them, RNAPIII has the most complex structure, consisting of 17 subunits, and it is primarily responsible for the synthesis of transfer RNA (tRNA). Compared with RNAPII consisting of 12 subunits, RNAPIII contains a unique heterotrimer Rpc82/31/34 and a heterodimer Rpc53/37 which is homologous to the counterpart of RNAPI. This paper reviews the structures and functions of the specific heterotrimer and heterodimer in RNAPIII, aiming to lay a theoretical foundation for further studies on the modification mechanisms and assembly processes of specific subunits of RNAPIII in S. cerevisiae.

Saccharomyces cerevisiae  /  RNA polymerase III  /  specific subunit  /  transcription
杨欣怡, 李盼, 曾凡力. 酿酒酵母RNA聚合酶特异亚基的研究进展. 微生物学报, 2025 , 65 (7) : 2854 -2863 . DOI: 10.13343/j.cnki.wsxb.20250006
Xinyi YANG, Pan LI, Fanli ZENG. Research progress in specific subunits of RNA polymerase III in Saccharomyces cerevisiae[J]. Acta Microbiologica Sinica, 2025 , 65 (7) : 2854 -2863 . DOI: 10.13343/j.cnki.wsxb.20250006
RNA聚合酶(RNA polymerase, RNAP)在所有真核生物中高度保守。它是一个由多亚基组成的复合体,负责催化生物体内的转录过程。在原核生物中,仅存在一种RNA聚合酶,而1969年Roeder等[1]发现真核生物中存在3种不同的RNA聚合酶,分别是RNA聚合酶I (RNAPI)、RNA聚合酶II (RNAPII)和RNA聚合酶III (RNAPIII)[1]。随后,研究者又发现了另外4种不同的真核RNA聚合酶,其中包括位于细胞质中的线粒体RNA聚合酶(mitochondrial RNA polymerase, POLRMT);以及3种仅存在于植物中的RNA聚合酶,其中2种聚合酶(RNAPIV和RNAPV)位于细胞核内,另有一种质体编码的RNA聚合酶(plastid-encoded RNA polymerase, PEP)仅存在于叶绿体中[2-10]
真核生物中主要存在RNAPI、RNAPII和RNAPIII等3种RNA聚合酶(图1)。RNAPI包含14个亚基;RNAPII包含12个亚基;RNAPIII结构最为复杂,包含17个亚基,其中包括一种独特的异三聚体Rpc82/34/31,以及一组异二聚体Rpc53/37,异二聚体在RNAPI中的同源物是Rpa34.5/49[11]。3种RNA聚合酶共有Rpb5、Rpb6、Rpb8、Rpb10和Rpb12等5个通用亚基[11-12];每种RNA聚合酶都有其独特的亚基,其中一些亚基能够在其他2种RNA聚合酶中找到同源物,例如在RNAPII中除了5个通用亚基外,其余7个亚基在RNA聚合酶I和III中都有相应的同源物;而有些亚基是独有的,例如RNAPIII中的Rpc82、Rpc34、Rpc31这3个亚基,在另外2种RNA聚合酶中无同源物[13]。这些独特的亚基赋予RNAPIII相应的独特性质和功能。
真核生物的RNAPIII负责转录所有转运RNA (transfer RNA, tRNA)、5S rRNA以及一些小型非编码RNA,约占细胞总RNA的15%。RNAPIII由17个亚基组成,其独有的异三聚体Rpc82/31/34在研究中被发现行使转录起始因子的功能[14],另外还有一对异二聚体Rpc53/37,它们参与转录的各个阶段,尤其在转录终止阶段具有重要作用。本文综述了RNAPIII的异三聚体和异二聚体的功能,以期为其亚基修饰与组装机制提供理论基础。
Rpc82/31/34是RNAPIII中独特的异三聚体,负责在RNAPIII中充当转录起始因子,在RNAPI和RNAPII中无对应的同源物[14]。细胞内转录的起始由转录因子与基因启动子结合开始,形成预起始复合物(preinitiation complex, PIC)。真核生物中的3种RNA聚合酶各自依赖一组特定的转录起始因子。RNAPII依赖多个起始因子启动转录,包括TATA结合蛋白(TATA-binding protein, Tbp)、TFIIA、TFIIB、TFIID、TFIIE、TFIIF和TFIIH。RNAPI利用Rrn3、Tbp、核心因子(core factor, CF)和上游相关因子(upstream activating factor, UAF)起始转录[15]
多位学者对RNAPIII的转录起始过程进行了深入探究。RNAPIII的转录起始首先由转录因子TFIIIA (仅用于5S rRNA转录)和TFIIIC识别DNA上的内部启动子元件(Box1和Box2);然后,TFIIIB (由Tbp、Brf1或称TFIIIIB70以及Bdp1或称TFIIIB90组成)被招募到被转录基因的上游区域,Brf1和Tbp协同组装到转录起始位点上游的DNA序列上,而Bdp1则主要通过其SANT结构域与Brf1-Tbp-DNA复合物结合[16-22];最后,TFIIIB将RNAPIII招募到转录起始位点,完成预起始复合物的形成[23] (图2)。已发现Brf1和Bdp1均与RNAPIII相互作用,并在启动子解开过程中发挥作用[24-27]。Tbp是RNA聚合酶I和II预起始复合物的通用亚基,Brf1与RNA聚合酶II的TFIIB转录起始因子相似,而Bdp1和TFIIIC是RNA聚合酶III的完全特异性转录起始因子[28-30]。转录因子在转录启动中起着关键作用,启动过程需要解开DNA双螺旋并进行初始的RNA合成。形成的DNA-RNA-RNAP三元复合体具有较高的稳定性[31]。生化数据表明,所有RNAP对RNA-DNA复合物具有高亲和力[32],RNAPIII中特异的亚基导致启动子识别和基因调控的差异。基于Rpc82和Rpc34的多个翼螺旋结构域(winged-helix domain, WHD)与RNAPII转录起始机制中的2个TFIIE亚基结构域相似,Rpc82/34/31三聚体被认为是一种TFIIE相关的亚复合体[33-34]
Rpc34是RNAPIII异三聚体的一个亚基,在PIC中与DNA启动子最上游的交叉连接中发挥重要作用[35]。Rpc34缺失或无义突变会导致细胞死亡[35]。Rpc34突变会减弱其与Brf1的相互作用[36];在酿酒酵母中,Rpc34缺陷型突变体的细胞中,tRNA合成减少,但5S rRNA合成未受影响[37],这表明Rpc34参与了RNAPIII的招募,并且仅作用于TFIIIB和TFIIIC之间的相互作用。研究发现2种Rpc34的突变体Rpc34-1124和Rpc34-1109在转录过程中对碱基招募与野生型Rpc34相比无差异,但Rpc34-1124突变体导致的转录缺陷可以通过增加底物中RNAPIII的量来补偿,表明该突变体影响的是RNAPIII与PIC的亲和性;而Rpc34-1109突变体在增加RNAPIII后并未恢复转录水平,说明该突变体是由于PIC形成受损而导致转录起始的缺陷[37]。此外,通过定点自由羟基探测揭示了Brf1细胞周期蛋白重复序列与Rpc34高度保守区域之间的结合,进一步佐证了Rpc34与Brf1之间的相互作用;Brf1的C端结构域还包含与Tbp的结合位点[38],推测Rpc34与Tbp之间可能存在未知的相互作用。对Brf1的3个保守基序进行突变,发现一些突变改变了与Rpc34亚基的相互作用,并且Brf1与Rpc34同时突变会导致细胞死亡,进一步佐证了预起始复合物TFIIIB-TFIIIC-DNA对RNAPIII的招募,部分通过RNAPIII的Rpc34亚基和Brf1之间的相互作用介导[39]
Rpc31是异三聚体中的另一个亚基,其突变会影响转录起始,但不会影响酶的普遍催化性质[40]。Rpc31在蛋白质精氨酸甲基转移酶1 (protein arginine methyltransferase 1, Hmt1)介导下发生甲基化,在tRNA基因转录中发挥2种作用:在最适合生长的条件下,甲基化的Rpc31正向调节tRNA基因的转录;在压力胁迫环境中,Rpc31与RNA聚合酶III抑制因子Maf1相互作用,进而抑制转录[41]。Maf1是RNAPIII的负调节因子,RNAPIII介导的转录在最佳环境下是稳定的,而在不利的生长条件下或遇到其他形式的胁迫后,RNAPIII的转录被Maf1抑制[42];具体机制涉及Maf1的磷酸化:压力条件下,Maf1被去磷酸化,然后从细胞质转移到细胞核,在细胞核中Maf1结合并调节RNAPIII的转录活性[43-44]。基于冷冻电子显微镜(Cryo-electron microscopy, Cryo-EM)技术对RNAPIII-Maf1复合物结构进行分析,揭示Maf1的结合会导致Rpc82/34/31异三聚体的重排[45],甲基化的Rpc31与Maf1相互作用,抑制Rpc34和TFIIIB亚基Brf1之间的相互作用,从而阻止RNAPIII被招募到启动子,表明Rpc31影响PIC的形成,而是否对后续转录过程进行调控仍不明确。
RNAPIII的结构分析表明,Rpc31的N末端区域锚定在Rpc82上,并进一步与RNA聚合酶III的核心和茎亚复合物相互作用[46];C末端结构域与RNAPIII的转录起始密不可分:Rpc31的C末端高度保守并富含Asp-Glu酸性氨基酸[47],通过定点光交联法绘制PIC内的蛋白质相互作用,揭示了这个酸性区域在转录起始期间特异性地靶向Rpc34,但也与游离的RNAPIII中的DNA相互作用,表明该区域在RNAPIII特异性转录起始因子相互作用中发挥作用[46]
Rpc82的正确修饰对RNAPIII全酶的组装和转录至关重要。细胞对营养物质的反应由雷帕霉素复合物1 (target of rapamycin complex 1, TORC1)的保守靶点调节[48-49]。对于RNAPIII,TORC1通过保守的RNAPIII抑制因子Maf1的磷酸化调节tRNA合成。在营养丰富的细胞环境中,Maf1会发生磷酸化修饰,而这一磷酸化过程由TORC1介导。经磷酸化修饰后的Maf1,其分子构象发生改变,导致无法进入细胞核,进而不能参与对RNAPIII转录过程的调控。当细胞面临氮耗尽的情况,或者添加特定的TORC1抑制剂雷帕霉素时,TORC1的活性会受到抑制,从而引发细胞的压力胁迫反应。在此情况下,Maf1不再被TORC1磷酸化,去磷酸化的Maf1能够与RNAPIII相互作用,并对RNAPIII的转录活性产生抑制作用,最终致使tRNA合成水平下调[50]。类泛素化是近年来基因修饰的研究热点,参与了多种细胞过程,包括信号传递、细胞核质运输、细胞周期进程、DNA损伤反应和转录[51-53]。研究发现,RNAPIII的许多成分在正常生长条件下都被类泛素化,例如亚基Rpc82、Rpc128和Rpc53。这种修饰不仅存在于酿酒酵母中,在人类细胞中也被频繁发现,说明其进化保守[54]。雷帕霉素处理会导致这些亚基的类泛素化剧烈减少,表明正确的类泛素化需要TORC1活性;将Rpc82中潜在的类泛素化位点K403、K406、K591和K596的赖氨酸替换为精氨酸,完全消除了Rpc82的类泛素化,后者的表达量未改变,但与野生型Rpc82细胞相比,tRNA基因的转录减少了约50%[55]。免疫共沉淀结果表明,类泛素化促进了Rpc82加入RNAPIII全酶[56],但目前的研究并未确切说明类泛素化是否介导了Rpc82向全酶的组装以及如何调控这一过程。此外,TORC1如何调节Rpc82的类泛素化仍不明确。
此外,研究发现Rpc82在多个残基上被磷酸化,其中一些磷酸化依赖于TORC1活性[57-58]。S392和S394位点似乎对雷帕霉素特别敏感,这2个位点位于蛋白激酶A (protein kinase A, PKA)互作基序中,表明它们被PKA以TORC1依赖性方式磷酸化;T430位于潜在的酪蛋白激酶2 (casein kinase 2, CK2)磷酸化位点,由于这些位点未经过氨基酸替代研究验证,因此尚不清楚S392、S394和T430的磷酸化如何影响Rpc82活性[56]。然而,鉴于TORC1、PKA和CK2促进RNAPIII活性,这些磷酸化事件很可能与RNAPIII活性的增加相关[56]。总结来说,Rpc82的类泛素化和磷酸化对于RNAPIII全酶的组装以及Rpc82招募到tRNA基因至关重要,Rpc82的TORC1依赖性类泛素化在最佳生长条件下增强了RNAPIII的转录能力。
RNAPIII的异二聚体Rpc53/37与其正常行使转录功能密切相关。Rpc53主要与转录因子相关联,而Rpc37则与转录终止的顺利进行密不可分[59]。RNA聚合酶转录起始因子与启动子DNA元件结合,招募RNA聚合酶,形成稳定的转录起始复合物,直到约10个核苷酸的新生RNA在活性中心与DNA形成RNA-DNA杂交链[60-61],并成为延伸复合物(elongation complex, EC)稳定性的主要组成部分,该复合物能够长时间持续合成RNA[62-63]。在转录结束时,必须破坏EC的稳定性,才能释放其中的RNA和DNA。与转录启动类似,EC的解体需要顺式信号和蛋白质因子介导,转录物才能从RNA聚合酶的抓持中释放[64]
三种聚合酶存在不同的转录终止策略。RNAPII催化的转录终止依赖于大多数真核生物基因末端包含的一段AATAAA共同序列,以及下游的一段富含GT的序列,这些序列被称为转录终止的修饰点[59];终止与mRNA的3′末端加工及多聚腺苷酸化紧密耦合,涉及3′末端加工活性与RNAPII最大亚基的羧基末端结构域之间的调节相互作用[65-67]。此外,RNAPII的转录终止可能需要终止区域的染色质重塑[68]。虽然驱动RNAPII转录终止的完整分子机制仍未完全阐明,但可以确定转录终止的第一步是聚合酶正确停滞在终止子区域[69]。RNAPI的终止机制涉及正确终止及释放转录物所需的顺式作用元件和反式作用因子[70]。RNAPI的终止至少需要序列特异性的DNA结合蛋白(如TTF1)充当阻拦聚合酶延伸的路障,并包括暂停位点上游的释放元件[71]。正常情况下,RNAPI的转录终止发生在终止因子DNA结合位点上游约10-12个碱基处。在小鼠和酵母中,上游元素与细胞因子或释放因子协同作用,细胞因子或释放因子与RNAPI相互作用并驱动暂停的RNA-DNA复合物的解体[72]。RNAPIII的终止信号与RNAPI和RNAPII不同。RNAPIII能够准确有效地识别由一小段胸腺嘧啶核苷残基构成的终止信号,并在尿嘧啶核苷延伸处暂停转录[73-74]
Rpc37在RNAPIII转录终止中发挥重要作用。酿酒酵母RNPIII的核心部分可以在缺失Rpc53/37亚基的情况下发挥作用,但其转录的终止需要合成8个或更多个3′端尿嘧啶核苷;全酶转录终止机制需要较少的尿苷,表明RNAPIII使用Rpc53/37来减缓延伸并使终止子顺利作用[75]。酿酒酵母RNAPIII终止前复合物(pretermination complex, PTC)的分离检测表明,Rpc53/37亚基是PTC形成所必需的,PTC的形成由终止子非模板链的近端部分引导;然而,PTC的形成并不能确保转录物的释放,因为终止子的模板链是EC解体的主要效应物,而非模板链携带通过Rpc53/37亚基发挥作用的不同序列,从而特异性发出终止信号[59]。Rpc37对于非模板链终止信号的这一功能至关重要,其C末端的部分缺失导致终止子区转录物释放的缺陷;随机突变的Rpc37突变体表现出明显的表型,表现为突变体细胞在转录时存在高达40%的通读,而且突变体从终止子中释放的新生前体tRNA的3′-oligoU片段更短,表明终止过程中tRNA的3′端发生断裂[76]
RNAPIII异二聚体Rpc53/37复合物还参与启动子打开以及转录起始[77]。异二聚体Rpc53/37的结构类似于TFIIF (RNAPII的转录起始因子)。因为异二聚体Rpc53/37显示出与TFIIF的α亚基和β亚基(Rap74和Rap30)相似的结构域,并且异二聚体Rpc53/37具有一个位于RNAPIII裂叶结构域中的保守二聚化模块,与TFIIF相似[78]。因此,推测Rpc53/37在RNAPIII的转录中也起到类似转录起始因子的作用,但目前缺少更直接的证据来支持这一观点。Rpc53的C末端区域与Rpc37二聚化,锚定在RNAPIII裂缝的叶片上[77],其中一个片段定位于RNAPIII延伸复合物的RNA-3′末端附近;Rpc53的N末端片段与另一个RNAPIII特异性亚复合体的Rpc82/Rpc34亚基以及转录起始因子TFIIIC相互作用;此外,还确定了Rpc37的C末端区域的一个重要片段位于RNAPIII活性中心的几个高度保守基序附近,并与转录因子TFIIIB的Bdp1亚基相互作用,表明Rpc53/37复合体也参与了转录起始[28]。将Rpc53的N末端进行定点丙氨酸替代突变,产生了冷敏感性生长缺陷菌株和RNAPIII体外转录活性受损菌株;其N末端高度无序的57个氨基酸链是一种通用的蛋白质结合模块,能够与Rpc37及转录起始因子TFIIIC的Tfc4亚基显示出高水平的结合亲和力[77]。与Rpc82类似,Rpc53的磷酸化也受TORC1调节,雷帕霉素处理后Rpc53的磷酸化和Maf1的去磷酸化共同抑制了RNA聚合酶III转录活性[79]。以上证据表明Rpc53/37异二聚体在RNAPIII介导的转录起始和终止中发挥重要作用。
RNA聚合酶III作为真核生物3种RNA聚合酶中最为复杂的一种,长期以来研究者们已对其特异性亚基的结构和功能进行了日益深入的研究。独特的异三聚体Rpc82/34/31在RNAPIII介导的转录过程中首先与转录起始因子接触,其结构与起始因子TFIIE相似,显示出该异三聚体在转录起始复合物形成中的关键作用。异二聚体Rpc53/37的功能更为复杂,既在转录起始时与起始因子保持高度相互作用,又在转录终止时协助RNAPIII高效、快速地完成停止。然而,关于这些过程涉及的亚基修饰机制的研究,诸如Rpc82和Rpc53的磷酸化及类泛素化,尤其是它们如何巧妙地调控自身与RNAPIII全酶的配合,以及如何在外界环境变化时精准有效地调节RNAPIII活性,仍是一个亟待深入探讨的领域。
此外,真核生物的RNA聚合酶通过多个亚基的组装形成具有转录活性的全酶,这一过程需要组装因子的参与。近年来,Liu等[80]的研究发现,Npa3/Gpn1分别与Gpn3和Rba50相互作用,并与RNAPII的第二大亚基Rpb2发生相互作用。随后,研究确定了Npa3和Gpn3直接参与了RNAPII两个大亚基的组装过程[81],且这一过程受Rtr1的协调参与[82]。此外,Rba50与Gpn2相互作用,协同组装RNAPII中的Rpb3亚复合体,随后发现这2个组装因子还促进了RNAPII与RNAPIII第二大亚基的装配[83-84]。然而,这些组装因子如何协同调控RNAPIII全酶组装的分子机制尚不明确,特别是特异性亚基装配到全酶所需的组装因子仍然未知。本综述以RNAPIII特异性亚基组成的异三聚体和异二聚体为切入点,通过研究其结构和功能,寻找相应的组装因子并探讨组装过程,为进一步研究RNA聚合酶III特异性亚基修饰机制及其组装提供理论基础。
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。
  • 河北省杰出青年自然科学基金(C2023204155)
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2025年第65卷第7期
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doi: 10.13343/j.cnki.wsxb.20250006
  • 接收时间:2025-01-02
  • 首发时间:2026-02-06
  • 出版时间:2025-07-04
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  • 收稿日期:2025-01-02
  • 录用日期:2025-02-24
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Natural Science Foundation of Hebei Province for Outstanding Youth(C2023204155)
河北省杰出青年自然科学基金(C2023204155)
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    河北农业大学 生命科学学院,河北 保定
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2种不同金属材料的力学参数

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种数
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species
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鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
小菇科 Mycenaceae 2 12 5.74 丝盖伞属 Inocybe 5 2.39
多孔菌科 Polyporaceae 8 14 6.70 蜡蘑属 Laccaria 5 2.39
红菇科 Russulaceae 3 23 11.00 小皮伞属 Marasmius 6 2.87
小菇属 Mycena 11 5.26
光柄菇属 Pluteus 5 2.39
红菇属 Russula 17 8.13
栓菌属 Trametes 5 2.39
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