BMP signaling can be fine-tuned through phosphorylation of Smad1/5/8 linker regions. Smad1 linker phosphorylation integrates diverse signals to control BMP signaling through interplay between phosphorylation, ubiquitination, and exclusion from the nucleus. The level of BMP activity in developing tissues is also fine-tuned by secreted proteins that modulate BMP interaction with receptors and proteases that regulate BMP trap proteins. The 1942 Spemann Organizer graft experiment demonstrated that transplanting the dorsal lip of a salamander embryo induced neighboring cells to adopt more dorsal fates, organizing nervous system, somite and kidney tissues and establishing patterning of the embryo.