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Calcium sensors of ciliary outer arm dynein: functions and phylogenetic considerations for eukaryotic evolution

机译：睫状外臂动力蛋白的钙传感器：真核生物进化的功能和系统发育考虑

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The motility of eukaryotic cilia and flagella is modulated in response to several extracellular stimuli. Ca2+ is the most critical intracellular factor for these changes in motility, directly acting on the axonemes and altering flagellar asymmetry. Calaxin is an opisthokont-specific neuronal calcium sensor protein first described in the sperm of the ascidian Ciona intestinalis. It binds to a heavy chain of two-headed outer arm dynein in a Ca2+-dependent manner and regulates ‘asymmetric’ wave propagation at high concentrations of Ca2+. A Ca2+-binding subunit of outer arm dynein in Chlamydomonas reinhardtii, the light chain 4 (LC4), which is a Ca2+-sensor phylogenetically different from calaxin, shows Ca2+-dependent binding to a heavy chain of three-headed outer arm dynein. However, LC4 appears to participate in ‘symmetric’ wave propagation at high concentrations of Ca2+. LC4-type dynein light chain is present in bikonts, except for some subclasses of the Excavata. Thus, flagellar asymmetry-symmetry conversion in response to Ca2+ concentration represents a ‘mirror image’ relationship between Ciona and Chlamydomonas. Phylogenetic analyses indicate the duplication, divergence, and loss of heavy chain and Ca2+-sensors of outer arm dynein among excavate species. These features imply a divergence point with respect to Ca2+-dependent regulation of outer arm dynein in cilia and flagella during the evolution of eukaryotic supergroups.
机译：真核纤毛和鞭毛的运动性受到几种细胞外刺激的调节。 Ca 2 + 是这些运动性变化的最关键的细胞内因子，直接作用于轴突并改变鞭毛不对称性。 Calaxin是视神经胶质蛋白特定的神经元钙传感器蛋白，首先在海鞘Ciona intestinalis的精子中描述。它以依赖Ca 2 + 的方式与两头外臂动力蛋白的重链结合，并在高浓度Ca 2 + 下调节“不对称”波的传播。莱茵衣藻中外臂动力蛋白的Ca 2 + 结合亚基，轻链4（LC4），是与钙调蛋白在系统发育上不同的Ca 2 + 传感器，显示Ca 2 + 依赖性结合到三头外臂动力蛋白的重链上。然而，LC4似乎在高浓度的Ca 2 + 中参与“对称”波传播。 LC4型达因轻链存在于bikonts中，除了Excavata的某些亚类外。因此，响应Ca 2 + 浓度的鞭毛不对称-对称转换代表了Ciona和衣藻之间的“镜像”关系。系统发育分析表明，挖掘物种中外臂动力蛋白的重链和Ca 2 + 传感器重复，发散和丢失。这些特征暗示了在真核超群进化过程中纤毛和鞭毛中Ca 2 + 依赖的外臂动力蛋白调控的分歧点。

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