Department of Neuroscience, Yale School of Medicine, 295 Congress Ave, New Haven, CT 06510, USA.
Genetic Models of Disease Laboratory, Oklahoma Medical Research Foundation, 825 N. E. 13th St, Oklahoma City, OK 73104, USA.
Curr Biol. 2023 Sep 25;33(18):3851-3864.e7. doi: 10.1016/j.cub.2023.07.052. Epub 2023 Aug 15.
Axonal transport is key to neuronal function. Efficient transport requires specific motor-cargo association in the soma, yet the mechanisms regulating this early step remain poorly understood. We found that EBP-1, the C. elegans ortholog of the canonical-microtubule-end-binding protein EB1, promotes the specific association between kinesin-3/KIF1A/UNC-104 and dense core vesicles (DCVs) prior to their axonal delivery. Using single-neuron, in vivo labeling of endogenous cargo and EBs, we observed reduced axonal abundance and reduced secretion of DCV cargo, but not other KIF1A/UNC-104 cargoes, in ebp-1 mutants. This reduction could be traced back to fewer exit events from the cell body, where EBP-1 colocalized with the DCV sorting machinery at the trans Golgi, suggesting that this is the site of EBP-1 function. EBP-1 calponin homology (CH) domain was required for directing microtubule growth on the Golgi, and mammalian EB1 interacted with KIF1A in an EBH-domain-dependent manner. Loss- and gain-of-function experiments suggest a model in which both kinesin-3 binding and guidance of microtubule growth at the trans Golgi by EBP-1 promote motor-cargo association at sites of DCV biogenesis. In support of this model, tethering either EBP-1 or a kinesin-3/KIF1A/UNC-104-interacting domain from an unrelated protein to the Golgi restored the axonal abundance of DCV proteins in ebp-1 mutants. These results uncover an unexpected role for a microtubule-associated protein and provide insights into how specific kinesin-3 cargo is delivered to the axon.
轴突运输对于神经元功能至关重要。有效的运输需要在体部进行特定的运动货物结合,但调节这一早期步骤的机制仍知之甚少。我们发现,C. elegans 中经典微管末端结合蛋白 EB1 的同源物 EBP-1,在其轴突运输之前,促进了驱动蛋白-3/KIF1A/UNC-104 与致密核心囊泡(DCV)之间的特异性结合。使用单个神经元、内源性货物和 EBs 的体内标记,我们观察到 ebp-1 突变体中 DCV 货物的轴突丰度降低和分泌减少,但其他 KIF1A/UNC-104 货物没有减少。这种减少可以追溯到从细胞体中出口事件减少,EBP-1 与 DCV 分拣机制在反高尔基体内共定位,这表明这是 EBP-1 功能的位点。EBP-1 钙调蛋白同源(CH)结构域对于指导高尔基体内微管的生长是必需的,并且哺乳动物 EB1 以 EBH 结构域依赖性的方式与 KIF1A 相互作用。缺失和功能获得实验表明了一种模型,即 EBP-1 既促进了驱动蛋白-3 的结合,又指导了反高尔基体内微管的生长,从而促进了 DCV 发生部位的运动货物结合。为了支持这个模型,将 EBP-1 或与 kinesin-3/KIF1A/UNC-104 相互作用的结构域从一种不相关的蛋白质中连接到高尔基体内,就可以恢复 ebp-1 突变体中 DCV 蛋白的轴突丰度。这些结果揭示了一个微管相关蛋白的意外作用,并提供了对特定驱动蛋白-3 货物如何递送到轴突的深入了解。