The BMP ligand Dpp, operates as a long range morphogen to control many important functions during Drosophila development from tissue patterning to growth. The BMP signal is transduced intracellularly via C-terminal phosphorylation of the BMP transcription factor Mad, which forms an activity gradient in developing embryonic tissues. Here we show that Cyclin dependent kinase 8 and Shaggy phosphorylate three Mad linker serines. We demonstrate that linker phosphorylations control the peak intensity and range of the BMP signal across rapidly developing embryonic tissues. Shaggy knockdown broadened the range of the BMP-activity gradient and increased high threshold target gene expression in the early embryo, while expression of a Mad linker mutant in the wing disc resulted in enhanced levels of C-terminally phosphorylated Mad, a 30% increase in wing tissue, and elevated BMP target genes. In conclusion, our results describe how Mad linker phosphorylations work to control the peak intensity and range of the BMP signal in rapidly developing Drosophila tissues.
Throughout the development of battery technologies in recent years, the solid-state electrolyte (SSE) has demonstrated outstanding advantages in tackling the safety shortcomings of traditional batteries while meeting high demands on electrochemical performances. The traditional manufacturing strategies can achieve the fabrication of batteries with simple forms (coin, cylindrical, and pouch), but encounter limitations in preparing complex-shaped or micro/nanoscaled batteries especially for inorganic solid electrolytes (ISEs). The advancement in novel manufacturing techniques like 3D printing has enabled the assembly of different solid electrolytes (polymeric, inorganic, and composites) in a more complex geometric configuration. However, there is a huge gap between the capabilities of the current 3D printing techniques and the requirements for battery production. In this review, we compare the traditional manufacturing to several novel 3D printing techniques, highlighting the potential of 3D printing in the SSE manufacturing. The latest SSE manufacturing progress in the group of direct-writing (DW) based or lithography-based printing technologies are summarized separately from the perspectives of feedstock selection, build envelope, printing resolution, and application (nano-scaled, flexible, and large-scale battery grids). Throughout the discussion, some challenges associated with manufacturing SSEs via 3D printing such as air/moisture sensitivity of samples, printing resolution, scale-up capability, and longterm sintering for ISEs have been put forward. This review aims to bridge the gap between 3D printing techniques and battery requirements by analyzing the existing limitation in SSE manufacturing and point out future needs.
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