Chapter 15. Neural Tube
Neural Tube: Birth of the Central Nervous System
Development of the Ectoderm
- BMP Levels and Ectoderm Fate
- Bone Morphogenetic Proteins (BMP) regulate ectoderm development.
- Low BMP levels: Ectoderm differentiates into the neural tube.
- Presence of BMP Inhibitors: Critical for neural tube formation.
Neurulation
- Definition of Neurulation: The process of forming the neural tube, crucial for central nervous system development.
- Cell Movement and Shape Change: Understanding how cells rearrange to create the neural structures.
Types of Neurulation
- Primary Neurulation:
- The neural plate rolls up to form a tube.
- Secondary Neurulation:
- Involves the formation of a solid mass of tissue that hollows out to create the neural tube.
- In mammals, starts at the sacral vertebrae region.
Neural Plate Formation
- The neural plate forms through signaling from the dorsal mesoderm, transforming ectoderm into columnar cells, forming a raised structure.
- Can include up to 50% of the ectoderm in its formation.
Tissue Shape Changes in Neurulation
- The neural plate becomes longer and narrower due to convergent extension, where cells move closer together.
- Cells also preferentially divide in the anterior-posterior direction rather than sideways, facilitating extension.
Neural Plate Bending
- Medial Hinge Point (MHP):
- Cells at the midline form MHP, which gets induced by the notochord to take a wedge shape.
- Dorsolateral Hinge Points:
- Found on either side, also forming wedge shapes, contributing to the tube formation by bending the neural plate.
- The surface ectoderm applies pressure on the neural plate, aiding in bending through sideways cell division.
Neural Tube Closure
- Neural Folds:
- They meet and adhere, leading to the formation of the neural tube.
- Cells at the edges form neural crest, which migrates to create peripheral neurons and other cell types.
- Regulation by Cadherins:
- Cells express E-cadherin initially.
- During neurulation, they begin to express N-cadherin to help separate neural tube from surface ectoderm, allowing neural crest cells to not express either cadherin.
Defects in Neural Tube Closure
- Approximately 1 in 1000 live births experience neural tube defects.
- Spina Bifida:
- Defect in posterior neural tube closure; may result in vertebrae not fusing or spinal cord bulging.
- Anencephaly:
- Defect in anterior neural tube closure leading to underdeveloped forebrain and skull.
- Folic Acid's Role:
- Folic acid deficiency correlated with neural tube defects; promotes proper DNA methylation.
- Supplements recommended for women of child-bearing age.
Cell-Cell Signaling: Dorsal-Ventral Differences
- Dorsal vs. Ventral:
- Dorsal side: Sensory neurons.
- Ventral side: Motor neurons.
- Signaling Molecules:
- BMP and TGF-β from ectoderm above, Sonic Hedgehog (Shh) from notochord below induce differentiation in the neural tube.
Gradient Formation
- TGF-β and Shh Gradients:
- Induction occurrences shaped by the notochord; TGF-β signals induce the dorsal region, while Shh induces the ventral region of the neural tube.
Consequences of Shh Expression
- Shh signaling suppresses Pax6 expression, important for eye formation.
- Abnormal Shh levels may cause defects, potentially resulting in cyclopia (one eye formation).