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The structural heterogeneity of radial spokes in cilia and flagella is conserved

机译:纤毛和鞭毛radial状辐条的结构异质性得以保留

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Radial spokes (RSs) are ubiquitous components of motile cilia and flagella and play an essential role in transmitting signals that regulate the activity of the dynein motors, and thus ciliary and flagellar motility. In some organisms, the 96 nm axonemal repeat unit contains only a pair of spokes, RS1 and RS2, while most organisms have spoke triplets with an additional spoke RS3. The spoke pairs in Chlamydomonas flagella have been well characterized, while spoke triplets have received less attention. Here, we used cryoelectron tomography and subtomogram averaging to visualize the three-dimensional structure of spoke triplets in Strongylocentrotus purpuratus (sea urchin) sperm flagella in unprecedented detail. Only small differences were observed between RS1 and RS2, but the structure of RS3 was surprisingly unique and structurally different from the other two spokes. We observed novel doublet specific features that connect RS2, RS3, and the nexin-dynein regulatory complex, three key ciliary and flagellar structures. The distribution of these doublet specific structures suggests that they could be important for establishing the asymmetry of dynein activity required for the oscillatory movement of cilia and flagella. Surprisingly, a comparison with other organisms demonstrated both that this considerable RS heterogeneity is conserved and that organisms with RS pairs contain the basal part of RS3. This conserved RS heterogeneity may also reflect functional differences between the spokes and their involvement in regulating ciliary and flagellar motility.
机译:径向辐条(RSs)是运动性纤毛和鞭毛的普遍组成部分,在传递调节达因蛋白运动的信号从而调节睫毛和鞭毛运动中起着至关重要的作用。在某些生物中,96 nm轴突重复单元仅包含一对辐条,即RS1和RS2,而大多数生物都具有三重峰,并带有附加的辐条RS3。衣原体鞭毛中的辐条对已被很好地表征,而辐条三胞胎受到的关注较少。在这里,我们使用了冷冻电子断层扫描和子断层图平均技术,以前所未有的细节可视化了紫斑海豚(Surpylocentrotus purpuratus)(海胆)精子鞭毛中轮辐三胞胎的三维结构。 RS1和RS2之间仅观察到很小的差异,但RS3的结构令人惊讶地独特,并且在结构上不同于其他两个辐条。我们观察到了连接RS2,RS3和nexin-dynein调节复合物,三个关键的睫毛和鞭毛结构的新型双峰特异特征。这些双态特异性结构的分布表明它们对于建立纤毛和鞭毛的振荡运动所需的动力蛋白的不对称性可能很重要。令人惊讶地,与其他生物的比较表明,这种相当大的RS异质性得到了保留,并且具有RS对的生物包含RS3的基础部分。这种保守的RS异质性也可能反映了辐条之间的功能差异及其在调节睫状和鞭毛运动中的作用。

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