Progress on the origin, function and evolutionary mechanism of RHR transcription factor family
Lv Menggang,1,2, Liu Aijia1,2, Li Qingwei,1,2, Su Peng,1,21 College of Life Science, Liaoning Normal University, Dalian 116081, China 2 Lamprey Research Center, Liaoning Normal University, Dalian 116081, China;
the National Natural Science Foundation of China.31801973 the National Natural Science Foundation of China.31772884 Dalian Supports Innovation and Entrepreneurship Projects of High Level Talents.2019RQ126 the Chinese Major State Basic Research Development Program (973).2013CB835304 the Marine Public Welfare Project of the State Oceanic Administration.201305016 the project of Department of Ocean and Fisheries of Liaoning Province.201805 and the Science and Technology Innovation Fund Research Project.2018J12SN079
NFAT(nuclear factor of activated T cells)蛋白首次在T细胞中被发现[29,30]。NFAT家族有5位成员分别为NFAT1(NFATc2或NFATp)、NFAT2(NFATc1)、NFAT3(NFATc4)、NFAT4(NFATc3)和NFAT5[31]。NFAT家族蛋白包含两个TAD结构域,一个调节结构域(NFAT homology region,NHR),一个高度保守的Rel结构域和一个羧基末端结构域[32]。NFAT家族分为两类,分别为经典家族和非经典家族。NFAT蛋白的经典的家族成员包括:NFAT1、NFAT2、NFAT3和NFAT4,具有钙调蛋白结合位点,受Ca2+和钙调蛋白的调控[33,34]。大量研究表明,钙调神经磷酸酶与经典的NFAT蛋白结合并使其脱磷酸化,从而诱导其进入细胞核并发挥转录活性,非经典家族成员仅包括NFAT5,其并不需要钙调神经磷酸酶参与其活性[35]。NFAT5可以形成同源二聚体,其同源二聚对与DNA结合和转录活性至关重要[36]。
RBP (recombination signal binding protein)家族蛋白对于神经系统的发育和造血系统的分化具有至关重要的作用[71],并且对于神经细胞分化的作用在生物进化过程中是保守的[72⇓⇓~75]。后生动物RBP蛋白有两种亚型,一种是RBP-Jκ (recombination signal binding protein for immunoglobulin kappa J region),其在Notch信号通路中发挥重要作用;另一种是RBP-L (recombination signal binding protein for immunoglobulin kappa J region like),其功能并不依赖于Notch信号通路[76]。RBP蛋白由3个结构域组成:N端结构域(N-terminal domain, NTD)、β折叠结构域(β-trefoil domain, BTD)和C端结构域(C-terminal domain, CTD)[77]。经过对其序列分析和结构研究证实,RBP的NTD与NF-κB转录因子家族的Rel同源结构域的N端具有相似性[78]。并且NTD以单体形式通过识别特异性序列(C/T)GTGGGAA与DNA结合,行使类似于Rel同源结构域的功能[79]。BTD结构域可以促进这种和DNA的结合,而且还能够参与蛋白质与蛋白质的相互作用,然而CTD并不与DNA发生相互作用[77,78]。除此之外,RBP分别通过与辅助激活蛋白和辅助抑制蛋白的不同相互作用,作为转录的激活因子和抑制因子发挥双重作用[80]。
A:EBF祖先基因中H1H2a序列的外显子-内含子的结构。0相位内含子(红色横线),1相位内含子(黑色横线)。B:H2复制的第一步是复制编码H2a的外显子。由于相位剪切规则,通过复制和选择性剪切产生了H2d的祖先基因。C:第二步是激活H2d编码区下游的隐蔽3′剪切位点(浅绿色三角)。D:产生了相同相位的i11和i12内含子,并将成熟的H2d嵌入EBF蛋白。根据文献[70]修改绘制。 Fig. 2Mechanisms of H2 replication in vertebrate EBF proteins
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