The head horn of the Asian rhinoceros beetle develops as extensively folded primordia before unfurling into its final 3D shape at the pupal molt. The information of the final 3D structure of the beetle horn is encoded in the folding pattern of the developing primordia. However, the developmental mechanism underlying epithelial folding of the primordia is unknown. In this study, we addressed this gap in our understanding of the developmental patterning of the 3D horn shape of beetles by focusing on the formation of surficial furrows that become the bifurcated 3D shape of the horn. By gene knockdown screening via RNAi, we found that knockdown of the gene Notch disturbed overall horn primordia furrow depth without affecting 2D furrow pattern. In contrast, knockdown of CyclinE altered 2D horn primordia furrow pattern without affecting furrow depth. From these results, depth and 2D pattern of primordial surficial furrow are likely to be regulated independently during the development and both of change can alter the final 3D shape.

中文翻译：

---

二维控制犀牛甲虫角3D形状的原始沟槽的深度和遗传控制

亚洲犀牛甲虫的头角发展为折叠的原基，然后在the蜕上展开成最终的3D形状。甲虫角的最终3D结构信息以显影原基的折叠模式编码。但是，原基上皮折叠的发育机制尚不清楚。在这项研究中，我们通过集中于表面沟槽的形成，解决了我们对甲虫3D角形状的发展模式的理解中的这一空白，这些表面成为角的分叉3D形状。通过RNAi进行的基因敲除筛选，我们发现Notch基因的敲除打乱了整个角原基犁沟深度，而没有影响2D犁沟模式。相反，敲除CyclinE可以改变2D角原基犁沟的犁纹，而不会影响犁沟的深度。