The Ectoderm: Neurulation, Neural Tube, Neural Crest
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Events that remove cells Progressive restriction from the epidermal of developmental lineage potential The ectoderm: neurulation, neural tube, neural crest Taube P. Rothman [email protected] Neural Shaping the neural plate induction 18 days PRIMARY NEURULATION Neural induction, Wnt Wnt formation of the neural plate Dorsal and Wnt Wnt ventral Formation of of the neural signaling in the groove and neural folds spinal cord Closure of neural Neural crest folds, formation of neural tube and neural crest Initially, the neural tube is composed of a single layer of neuroepithelial cells 1 Dorsal view: Neurulating embryos Dorsal view: Neurulating embryos REGIONS OF NEURAL TUBE CLOSURE Days 21-22 Day 23 cranioschisis meningomyecele REGIONALIZATION OF THE CNS PRIMARY AND SECONDARY VESICLES AND FLEXURES Cephalic Rhombomere flexure Cervical flexure NEUROGENESIS IN THE NEUROEPITHELIUM How are billions of CNS cells (neurons and glia) generated? lumen Cerebral cortex lumen Post mitotic daughter cell Lumen Nuclei (but not cells) migrate during the cell The plane of division of progenitor cells in the cycle ventricular zone influences their fate The neuroepithelium is a single layer of rapidly dividing stem cells 2 At least half the neurons that are generated undergo apoptosis and die. Neuronal survival depends upon recognition of target related trophic signals. Ectomesenchyme The neural crest Neural crest cells are guided through permissive Wnt BMP environments to RhoB reach their targets slug Ephrin proteins inhibit neural crest migration THE REGION OF THE NEURAXIS FROM WHICH A CREST CELL MIGRATES DETERMINES THE TARGET REACHED BY ITS DERIVATIVES cranial circumpharyngeal neural crest truncal cardiac vagal truncal sacral 3 Genetic potential, developmental restriction and differentiation •Some neural crest cells appear to be pluripotent. They can generate a remarkable number of differentiated cell types, but express only those phenotypes that are appropriate for the region they colonize. •Example: Cranial neural crest ectoderm: •The genetic potential of other crest-derived The fates of these crest derived cells cells is more restricted. There are a limited are controlled by the specific growth neurulation, neural tube, neural crest number of options in their genetic factors they encounter repertoire. The end •Example: truncal neural crest •Finally, some pre-migratory crest cells appear to be programmed for a specific developmental fate or if they are not committed to one before leaving the crest, there developmental options are severely Taube P. Rothman restricted during their migration. [email protected] •Example: cardiac neural crest 4.