Development

Cortical Radial Unit Hypothesis

  • The cortical radial unit hypothesis suggests that a radial column of the cortex comes from the same location in the ventricular zone.

  • Neurite & Symmetrical Cleavage:

    • Daughter cells exhibit different transcription factors (TFs).

    • The human cortex is significantly larger due to evolutionary factors.

  • Key Components:

    • Site radial Notch: Symmetrical division influenced by transcription factors including EMX2 and Pax6.

    • EMX2 is primarily involved in posterior development, while Pax6 is involved in anterior cortex development along the anterior-posterior (A-P) axis.

  • Neuronal Fate Decisions:

    • Neurons destined for the anterior cortex show higher levels of Pax6.

    • Notch signaling promotes the maintenance of progenitor cells in the posterior cortex.

    • Posterior cortex neurons exhibit higher levels of EMX2 and differentiate into astrocytes if EMX2 expression is low.

    • This leads to smaller posterior cortical areas and larger anterior regions.

    • Numb Protein:

    • Acts as a neural gate for fate determination during cleavage.

  • Pax6 Knockout Consequences:

    • Causal effect: Causal growth of posterior areas and shrinkage of anterior cortex.

    • Location in the ventricular zone influences thalamic organization and recruitment of thalamic neurons to corresponding subplate areas.

Neural Pathways Development

  • Pathway Selection:

    • Pathway selection involves making decisions along the proliferative zone, which is influenced by location, cell type, and cell-cell signaling.

    • Example: Origin of pyramidal neurotransmitters and astrocytes along the optic stalk shows three path choices:

    1. Enter optic tract on the same side.

    2. Enter the opposite side.

    3. Navigate through different optic nerves.

    • These decisions are influenced by extrinsic signals and local extracellular conditions along the subventricular zone.

  • Target Selection:

    • Determines the specific area of the brain to innervate.

    • Example: Selecting the correct thalamic nucleus, where neural precursor cells migrate along radial glial fibers to reach the lateral geniculate nucleus (LGN).

  • Address Selection:

    • Final decision on which part of the cortex or brain to innervate.

    • Example: Sorting and retraction of retinal axons aids in establishing retinotopy.

Neurite Growth Mechanisms

  • Growth Cones:

    • Neurite ends that identify targets using growth cues which help navigate towards synaptic targets.

  • Cellular Structures:

    • Lamellipodia: Flat sheets that undulate to explore the environment.

    • Filopodia: Projections that sample the local environment and grab substrate to elongate the axon.

  • Guidance Cues:

    • Permissive and Inhibitory Signals:

    • Neurites encounter a combination of permissive (growth-accommodating) and inhibitory signals that form guidance corridors.

    • CAMs (cell-adhesion molecules) help axons stick to the correct pathway.

  • Extracellular Signals:

    • Semaphorin 30:

    • A repulsive signal secreted by cells in the marginal zone that directs pyramidal axons downward while attracting dendrites.

Neural Development and Regeneration

  • Pioneer Axons:

    • Establish pathways for follower axons.

    • Example: In frogs, pioneer axons in the tectum receive inputs from the contralateral eye, using environmental cues for navigation.

  • Chemoattraction and Repulsion:

    • Netrins: Chemotactic signals that attract certain axons, facilitating growth into the anterior tectum.

    • Slit: A repulsive signal that guides axons by creating gradients leading to correct decussation.

  • Synapse Formation:

    • Neuromuscular junctions involve presynaptic and postsynaptic proteins that communicate neurotransmission.

    • Ephrin B:

    • Important in determining decussation patterns; secreted in gradients to modulate axonal pathways.

Neurogenesis and Neural Repair Mechanisms

  • Neurogenesis Discovery:

    • Joseph Altman in the 1980s initially described adult neurogenesis. Dr. Elisabeth discovered it in specific brain regions, namely the hippocampus and olfactory bulb.

  • Method to Identify Neurogenesis:

    • BrdU assay: A method to identify new neurons by incorporating bromodeoxyuridine (BrdU) during DNA synthesis in dividing cells.

  • Neurogenesis Process:

    • Precursor cells along the lateral ventricle migrate into designated areas (e.g., olfactory bulb) via the rostral migratory stream.

    • Some neurons in specific regions like the dentate gyrus of the hippocampus adopt GABA-ergic identities.

Cortical Radial Unit Hypothesis
  • The cortical radial unit hypothesis suggests that a radial column of the cortex comes from the same location in the ventricular zone.

  • Neurite & Symmetrical Cleavage:

    • Asymmetrical cleavage: Produces daughter cells with different fates and varying transcription factor profiles.

    • Daughter cells exhibit different transcription factors (TFs).

    • The human cortex is significantly larger due to evolutionary factors.

  • Key Components:

    • Site radial Notch: Symmetrical division influenced by transcription factors including EMX2 and Pax6.

    • EMX2 is primarily involved in posterior development, while Pax6 is involved in anterior cortex development along the anterior-posterior (A-P) axis.

  • Neuronal Fate Decisions:

    • Neurons destined for the anterior cortex show higher levels of Pax6.

    • Notch signaling promotes the maintenance of progenitor cells in the posterior cortex.

    • Posterior cortex neurons exhibit higher levels of EMX2 and differentiate into astrocytes if EMX2 expression is low.

    • This leads to smaller posterior cortical areas and larger anterior regions.

    • Numb Protein:

    • Acts as a neural gate for fate determination during cleavage.

  • Pax6 Knockout Consequences:

    • Causal effect: Causal growth of posterior areas and shrinkage of anterior cortex.

    • Location in the ventricular zone influences thalamic organization and recruitment of thalamic neurons to corresponding subplate areas.

Neural Pathways Development
  • Pathway Selection:

    • Pathway selection involves making decisions along the proliferative zone, which is influenced by location, cell type, and cell-cell signaling.

    • Example: Origin of pyramidal neurotransmitters and astrocytes along the optic stalk shows three path choices:

    1. Enter optic tract on the same side.

    2. Enter the opposite side.

    3. Navigate through different optic nerves.

    • These decisions are influenced by extrinsic signals and local extracellular conditions along the subventricular zone.

  • Target Selection:

    • Determines the specific area of the brain to innervate.

    • Example: Selecting the correct thalamic nucleus, where neural precursor cells migrate along radial glial fibers to reach the lateral geniculate nucleus (LGN).

  • Address Selection:

    • Final decision on which part of the cortex or brain to innervate.

    • Example: Sorting and retraction of retinal axons aids in establishing retinotopy.

Neurite Growth Mechanisms
  • Growth Cones:

    • Neurite ends that identify targets using growth cues which help navigate towards synaptic targets.

  • Cellular Structures:

    • Lamellipodia: Flat sheets that undulate to explore the environment.

    • Filopodia: Projections that sample the local environment and grab substrate to elongate the axon.

  • Guidance Cues:

    • Permissive and Inhibitory Signals:

    • Neurites encounter a combination of permissive (growth-accommodating) and inhibitory signals that form guidance corridors.

    • CAMs (cell-adhesion molecules) help axons stick to the correct pathway.

  • Extracellular Signals:

    • Semaphorin 30:

    • A repulsive signal secreted by cells in the marginal zone that directs pyramidal axons downward while attracting dendrites.

Neural Development and Regeneration
  • Pioneer Axons:

    • Establish pathways for follower axons.

    • Example: In frogs, pioneer axons in the tectum receive inputs from the contralateral eye, using environmental cues for navigation.

  • Chemoattraction and Repulsion:

    • Netrins: Chemotactic signals that attract certain axons, facilitating growth into the anterior tectum.

    • Slit: A repulsive signal that guides axons by creating gradients leading to correct decussation.

  • Synapse Formation:

    • Neuromuscular junctions involve presynaptic and postsynaptic proteins that communicate neurotransmission.

    • Ephrin B:

    • Important in determining decussation patterns; secreted in gradients to modulate axonal pathways.

Neurogenesis and Neural Repair Mechanisms
  • Neurogenesis Discovery:

    • Joseph Altman in the 1980s initially described adult neurogenesis. Dr. Elisabeth discovered it in specific brain regions, namely the hippocampus and olfactory bulb.

  • Method to Identify Neurogenesis:

    • BrdU assay: A method to identify new neurons by incorporating bromodeoxyuridine (BrdU) during DNA synthesis in dividing cells.

  • Neurogenesis Process:

    • Neural precursor cells along the lateral ventricle migrate into designated areas (e.g., olfactory bulb) via the rostral migratory stream.

    • Some neurons in specific regions like the dentate gyrus of the hippocampus adopt GABA-ergic identities.

    • Neurons complete differentiation: After migration and adopting identities, neurons mature and integrate into functional circuits.