Organizer induction requires cooperation between Siamois, Twin, and:
Formation of functional Spemann organizer requires integration of two vegetal signals: dorsal Wnt-beta-catenin via Siamois/Twin and Nodal-related TGF-beta via Smad2. Nodal ligands Xnr1,2,5 activated by VegT phosphorylate Smad2 which enters nucleus and cooperates with Siamois/Twin on enhancers of goosecoid, chordin and Cerberus. Chromatin immunoprecipitation shows Smad2/Siamois co-binding. Either pathway alone insufficient; blocking Smad2 with Cerberus-S or Siamois with dominant-negative abolishes organizer. Therefore cooperation between Siamois, Twin and Smad2 creates robust organizer inductio
Ref: Gilbert, Developmental Biology 12th ed., Chapter 10: Organizer induction by Siamois, Twin and Smad2 cooperation.