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Vertebrate Embryonic Patterning 3

Vertebrate Embryonic Patterning 3. Molecular Basis of Spemann’s Organizer. Inducing Centers in the Blastula. “Spemann”. (General). Fate Map of Organizer and Nieuwkoop’s Center. Molecular Nature of the Organizer. Transcription Factors Diffusible Molecules BMP inhibitors Wnt inhibitors.

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Vertebrate Embryonic Patterning 3

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  1. Vertebrate Embryonic Patterning 3 Molecular Basis of Spemann’s Organizer

  2. Inducing Centers in the Blastula “Spemann” (General)

  3. Fate Map of Organizer and Nieuwkoop’s Center

  4. Molecular Nature of the Organizer • Transcription Factors • Diffusible Molecules • BMP inhibitors • Wnt inhibitors

  5. Goosecoid • named based on homology to Drosophila gooseberry and bicoid homeodomain • expressed in the organizer – dorsal blastopore lip • activates txn of other organizer-specific genes e.g. Xotx2, Xnot, Xlim1, chordin, noggin, Xnr 1, 2, 4

  6. Goosecoid Expression Pattern Visit the Xenopus Molecular Marker Page

  7. Targeted Injections of Xenopus Embryos

  8. Goosecoid Induces Secondary Axes Injection of gsc into ventral vegetal blastomere at 8-16 cell stage induces a secondary axis. Both axes begin with the concentration of pigment at a blastopore lip (arrows). The 2° axis forms anterior head structures (cement gland)

  9. ORGANIZER MOLECULES BMP Inhibitors Noggin – expressed in deep mesoderm of organizer that ultimately forms notochord Chordin – expressed in deep mesoderm & pharyngeal endoderm that ultimately underlies head Cerberus – expressed in organizer & endoderm ultimately forming head

  10. BMP-4 Expression Pattern dorsal lip St. 10.5vegetal view vegetal view dorsal view St. 11

  11. Noggin Expression Pattern notochord D V

  12. Noggin Rescues UV Ventralized Embryos UV Noggin cloned through a functional screen by Richard Harland. Protein functions by binding to BMP2 & 4 and prevents their binding to their receptors. lo injected noggin mRNA hi

  13. Chordin Expression Pattern prechordal plate dorsal blastopore lip axial mesoderm blastopore St. 10 St. 10.5 St. 13

  14. Chordin Function Chordin also binds & sequesters BMP4

  15. BMP Gradient formed by Opposing Activities of Chordin & Xolloid

  16. Cerberus Expression Pattern

  17. Cerberus Duplicates Axes! Who’d a Thunk It?

  18. Cerberus Function • Cerberus binds to • Xwnt-8 (ventral mesoderm) • BMP-4 (ventral ectoderm) • Xnr-1 (dorsal mesoderm) Cerberus

  19. Cer Double Axes Enhanced by Blocking Wnt, BMP, and Xnr Pathways

  20. Follistatin • binds to TGF- family ligands • blocks their interaction with receptors

  21. Wnt Inhibitors • Frisbee – Soluble protein that mimics the frizzled wnt binding domain. • Dickkopf – functions by sequestering wnts • Cerberus – binds to wnts

  22. Frisbee Expression

  23. Frzb Function

  24. Frisbee Causes Anteriorization

  25. Frz & Fz are Members of Same Protein Superfamily Family CRD= cysteine rich domain: conserved set of cys residues that give protein distinct 2° structure

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