Hippo signaling is an important regulator of tissue size, but it also has a lesser-known role in tissue morphogenesis. Here we use the Drosophila pupal eye to explore the role of the Hippo effector Yki and its cofactor Mask in morphogenesis. We found that Mask is required for the correct distribution and accumulation of adherens junctions and appropriate organization of the cytoskeleton. Accordingly, disrupting mask expression led to severe mis-patterning and similar defects were observed when yki was reduced or in response to ectopic wts. Further, the patterning defects generated by reducing mask expression were modified by Hippo pathway activity. RNA-sequencing revealed a requirement for Mask for appropriate expression of numerous genes during eye morphogenesis. These included genes implicated in cell adhesion and cytoskeletal organization, a comprehensive set of genes that promote cell survival, and numerous signal transduction genes. To validate our transcriptome analyses, we then considered two loci that were modified by Mask activity: FER and Vinc, which have established roles in regulating adhesion. Modulating the expression of either locus modified mask mis-patterning and adhesion phenotypes. Further, expression of FER and Vinc was modified by Yki. It is well-established that the Hippo pathway is responsive to changes in cell adhesion and the cytoskeleton, but our data indicate that Hippo signaling also regulates these structures.

Hippo 信号传导是组织大小的重要调节器，但它在组织形态发生中也有一个鲜为人知的作用。在这里，我们使用果蝇蛹的眼睛来探索河马效应器 Yki 及其辅助因子面具在形态发生中的作用。我们发现，粘附连接的正确分布和积累以及细胞骨架的适当组织需要 Mask。因此，当yki减少或响应异位wts时，破坏掩码表达会导致严重的错误图案和类似的缺陷。此外，减少掩模产生的图案缺陷Hippo 通路活性改变了表达。RNA 测序揭示了在眼睛形态发生过程中适当表达大量基因需要 Mask。其中包括与细胞粘附和细胞骨架组织有关的基因、一组促进细胞存活的综合基因以及众多信号转导基因。为了验证我们的转录组分析，我们考虑了两个被 Mask 活性修饰的位点：FER和Vinc，它们在调节粘附方面发挥了作用。调节任一基因座修饰掩码错误图案和粘附表型的表达。此外，FER和Vinc 的表达由Yki修改。众所周知，Hippo 通路对细胞粘附和细胞骨架的变化有反应，但我们的数据表明，Hippo 信号也调节这些结构。