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Rapid Evolutionary Rewiring of a Structurally Constrained Eye Enhancer

机译：结构受限的眼部增强器的快速进化重新布线

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Background: Enhancers are genomic cis-regulatory sequences that integrate spatiotemporal signals to control gene expression. Enhancer activity depends on the combination of bound transcription factors as well as in some cases the arrangement and spacing of binding sites for these factors. Here, we examine evolutionary changes to the sequence and structure of sparkling, a Notch/EGFR/Runx-regulated enhancer that activates the dPax2 gene in cone cells of the developing Drosophila eye.Results: Despite functional and structural constraints on its sequence, sparkling has undergone major reorganization in its recent evolutionary history. Our data suggest that the relative strengths of the various regulatory inputs into sparkling change rapidly over evolutionary time, such that reduced input from some factors is compensated by increased input from different regulators. These gains and losses are at least partly responsible for the changes in enhancer structure that we observe. Furthermore, stereotypical spatial relationships between certain binding sites ("grammar elements") can be identified in all sparkling orthologs-although the sites themselves are often recently derived. We also find that low binding affinity for the Notch-regulated transcription factor Su(H), a conserved property of sparkling, is required to prevent ectopic responses to Notch in noncone cells.Conclusions: Rapid DNA sequence turnover does not imply either the absence of critical cis-regulatory information or the absence of structural rules. Our findings demonstrate that even a severely constrained cis-regulatory sequence can be significantly rewired over a short evolutionary timescale.
机译：背景：增强子是基因组顺式调控序列，整合时空信号以控制基因表达。增强子活性取决于结合的转录因子的结合以及在某些情况下这些因子的结合位点的排列和间隔。在这里，我们检查了Sparkling的序列和结构的进化变化，Notch / EGFR / Runx调节的增强子激活了果蝇眼发育中视锥细胞中的dPax2基因。在最近的进化历史中经历了重大的重组。我们的数据表明，各种监管投入到闪闪发光的相对强度会随着进化的时间而迅速变化，从而某些因素的减少的投入可以通过不同监管者的增加的投入得到补偿。这些收益和损失至少部分负责我们观察到的增强剂结构的变化。此外，尽管所有位点本身经常是最近才衍生出来的，但可以在所有波光粼粼的直系同源物中识别某些结合位点（“语法元素”）之间的定型空间关系。我们还发现，为了防止非锥细胞对Notch的异位应答，需要Notch调控的转录因子Su（H）的保守结合亲和力低，以防止异位细胞对Notch的异位反应。关键的顺式调控信息或缺乏结构规则。我们的发现表明，即使是严重受约束的顺式调控序列，也可以在较短的进化时间范围内显着重新连接。

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