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Anciently duplicated Broad Complex exons have distinct temporal functions during tissue morphogenesis

机译:古代复制的广泛复合物外显子在组织形态发生过程中具有独特的时间功能

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摘要

Broad Complex (BRC) is an essential ecdysone-pathway gene required for entry into and progression through metamorphosis in Drosophila melanogaster. Mutations of three BRC complementation groups cause numerous phenotypes, including a common suite of morphogenesis defects involving central nervous system (CNS), adult salivary glands (aSG), and male genitalia. These defects are phenocopied by the juvenile hormone mimic methoprene. Four BRC isoforms are produced by alternative splicing of a protein-binding BTB/POZ-encoding exon (BTB ( BRC )) to one of four tandemly duplicated, DNA-binding zinc-finger-encoding exons (Z1 ( BRC ), Z2 ( BRC ), Z3 ( BRC ), Z4 ( BRC )). Highly conserved orthologs of BTB ( BRC ) and all four Z ( BRC ) were found among published cDNA sequences or genome databases from Diptera, Lepidoptera, Hymenoptera, and Coleoptera, indicating that BRC arose and underwent internal exon duplication before the split of holometabolous orders. Tramtrack subfamily members, abrupt, tramtrack, fruitless, longitudinals lacking (lola), and CG31666 were characterized throughout Holometabola and used to root phylogenetic analyses of Z ( BRC ) exons, which revealed that the Z ( BRC ) clade includes Z ( abrupt ). All four Z ( BRC ) domains, including Z4 ( BRC ), which has no known essential function, are evolving in a manner consistent with selective constraint. We used transgenic rescue to explore how different BRC isoforms contribute to shared tissue-morphogenesis functions. As predicted from earlier studies, the common CNS and aSG phenotypes were rescued by BRC-Z1 in rbp mutants, BRC-Z2 in br mutants, and BRC-Z3 in 2Bc mutants. However, the isoforms are required at two different developmental stages, with BRC-Z2 and -Z3 required earlier than BRC-Z1. The sequential action of BRC isoforms indicates subfunctionalization of duplicated Z ( BRC ) exons even when they contribute to common developmental processes.
机译:广泛复合体(Broad Complex,BRC)是果蝇进入并通过变态发展所必需的蜕皮激素途径基因。三个BRC互补组的突变会导致许多表型,包括涉及中枢神经系统(CNS),成年唾液腺(aSG)和男性生殖器的常见形态发生缺陷。这些缺陷是由模拟荷尔蒙的幼年激素表型表现出来的。通过将蛋白结合的BTB / POZ编码外显子(BTB(BRC))与四个串联复制的,DNA结合的锌指编码外显子(Z1(BRC),Z2(BRC) ),Z3(BRC),Z4(BRC))。在双翅目,鳞翅目,膜翅目和鞘翅目的已发表cDNA序列或基因组数据库中发现了BTB(BRC)和所有4个Z(BRC)高度保守的直系同源物,表明BRC出现并在同源分裂之前进行了内部外显子复制。整个Holometabola都对电车轨道亚科成员,突变,电车轨道,无果,纵向缺失(lola)和CG31666进行了表征,并用于对Z(BRC)外显子进行系统发育分析,这表明Z(BRC)进化枝包括Z(突变)。没有已知基本功能的所有四个Z(BRC)域,包括Z4(BRC),都以与选择性约束一致的方式进化。我们使用转基因抢救来探索不同的BRC亚型如何促进共享的组织形态发生功能。如早期研究所预测,rbp突变体中的BRC-Z1,br突变体中的BRC-Z2和2Bc突变体中的BRC-Z3拯救了常见的CNS和aSG表型。但是,在两个不同的发育阶段都需要同工型,其中BRC-Z2和-Z3比BRC-Z1早。 BRC亚型的顺序作用表明重复的Z(BRC)外显子具有亚功能,即使它们有助于共同的发育过程。

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