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Context-specific requirements of functional domains of the Spectraplakin Short stop in vivo

机译：体内Spectraplakin功能域的上下文特定要求

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Spectraplakins are large multifunctional cytoskeletal interacting molecules implicated in various processes, including gastrulation, wound healing, skin blistering and neuronal degeneration. It has been speculated that the various functional domains and regions found in Spectraplakins are used in context-specific manners, a model which would provide a crucial explanation for the multifunctional nature of Spectraplakins. Here we tested this possibility by studying domain requirements of the Drosophila Spectraplakin Short stop (Shot) in three different cellular contexts in vivo: (1) neuronal growth, which requires dynamic actin-microtubule interaction; (2) formation and maintenance of tendon cells, which depends on highly stabilised arrays of actin filaments and microtubules, and (3) compartmentalisation in neurons, which is likely to involve cortical F-actin networks. Using these cellular contexts for rescue experiments with Shot deletion constructs in shot mutant background, a number of differential domain requirements were uncovered. First, binding of Shot to F-actin through the first Calponin domain is essential in neuronal contexts but dispensable in tendon cells. This finding is supported by our analyses of shot(kakP2) mutant embryos, which produce only endogenous isoforms lacking the first Calponin domain. Thus, our data demonstrate a functional relevance for these isoforms in vivo. Second, we provide the first functional role for the Plakin domain of Shot, which has a strong requirement for compartmentalisation in neurons and axonal growth, demonstrating that Plakin domains of long Spectraplakin isoforms are of functional relevance. Like the Calponin domain, also the Plakin domain is dispensable in tendon cells, and the currently assumed role of Shot as a linker of microtubules to the tendon cell surface may have to be reconsidered. Third, we demonstrate a function of Shot as an actin-microtubule linker in dendritic growth, thus shedding new light into principal growth mechanisms of this neurite type. Taken together, our data clearly support the view that Spectraplakins function in tissue-specific modes in vivo, and even domains believed to be crucial for Spectraplakin function can be dispensable in specific contexts.
机译：Spectraplakins是涉及多种过程的大型多功能细胞骨架相互作用分子，包括胃造胃，伤口愈合，皮肤起泡和神经元变性。据推测，Spectraplakins中发现的各种功能域和区域都以特定于上下文的方式使用，该模型将为Spectraplakins的多功能性质提供至关重要的解释。在这里，我们通过研究果蝇Spectraplakin短停点（Shot）在三种不同细胞背景下的体内结构域需求来测试这种可能性：（1）神经元生长，需要动态的肌动蛋白-微管相互作用; （2）肌腱细胞的形成和维持，这取决于肌动蛋白丝和微管的高度稳定的阵列，以及（3）神经元中的区室化，这可能涉及皮质F-肌动蛋白网络。使用这些细胞环境用于在shot突变体背景中使用Shot缺失构建体进行抢救实验，发现了许多不同的域要求。首先，Shot通过第一个Calponin域与F-肌动蛋白的结合在神经元环境中是必不可少的，但在肌腱细胞中却是必不可少的。这一发现得到了我们对shot（kakP2）突变体胚胎的分析的支持，而shot（kakP2）突变体胚胎仅产生缺少第一个钙蛋白结构域的内源同工型。因此，我们的数据证明了这些同工型在体内的功能相关性。其次，我们为Shot的Plakin域提供了第一个功能性角色，它对神经元和轴突生长的区隔有强烈的要求，这表明长Spectraplakin同工型的Plakin域具有功能相关性。像钙蛋白结构域一样，Plakin结构域在肌腱细胞中也是可有可无的，并且可能必须重新考虑目前假定的Shot作为微管与肌腱细胞表面的连接体的作用。第三，我们证明了Shot在树突状细胞生长中作为肌动蛋白-微管接头的功能，从而为这种神经突类型的主要生长机制提供了新的思路。两者合计，我们的数据清楚地支持这样的观点：Spectraplakins在体内以组织特异性模式起作用，甚至在某些情况下，甚至被认为对Spectraplakin功能至关重要的域也可以取消。

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