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Steps in setting up the Nervous system. Induction and Patterning of a neuron-forming region Birth and migration of neurons and glia Specification of cell fates Guidance of axonal growth cones to specific targets Formation of synaptic connections
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Steps in setting up the Nervous system • Induction and Patterning of a neuron-forming region • Birth and migration of neurons and glia • Specification of cell fates • Guidance of axonal growth cones to specific targets • Formation of synaptic connections • Binding of trophic factors for survival and differentiation • Competitive rearrangement of functional synapses • Continued synaptic plasticity during lifetime
Neuronal development • Ectoderm differentiation: Epidermis, nerve tissue, or neural crest cells? • Nerve tissue: Neuron or glial cell?
Neuronal development • Ectoderm differentiation: Epidermis, nerve tissue, or neural crest cells? • Nerve tissue: Neuron or glial cell?
Nerve Outgrowth • Filapodia-based movements • Rely on Cytoskeleton and factors that rearrange the cytoskeleton (GTPases).
Cerebellum • Important for motor control • Purkinje fibers • BMP and Granular layer
Movement of Neurons • Glial cells serve as track. • Nerve Birthdays: Determine how deep in the cortex a neuron will enter.
Cerebrum • Similar to Cerebellum • Neocortex: subdivided into 6 layers (old cells closest to ventricular zone). • Cortex divided horizontally (approx. 40 sections). • Fate decided early on (before S phase)
Motor Neuron Specification • Sonic hedgehog (Shh) secreted by floor plate.
Target specification of Motor neuron • As Nerves pass through the cortex, different transcriptional regulators are expressed. • Limb innervation determined by repulsion • If Lim1 is present, Eph A4 (receptor) is expressed; Eph A5 repels neuron • If no Lim1, neuropilin-2 expressed. (Axial muscles)
Following diffusible signals • Netrin-1 and Netrin-2 • SLIT and ROBO
Nerve Outgrowth • Filapodia-based movements • Rely on Cytoskeleton and factors that rearrange the cytoskeleton (GTPases).
Neural Crest Cells • Neurons/glial cells of sensory, sympathetic, and parasympathetic nerves • Epinephrine-producing cells of adrenal gland • Pigment containing cells • Skeletal/connective tissue of head • 4th germ layer?
Neural Crest Regionalization • Cranial • Cartilage, bone, cranial nerves, glia, connective tissues • Cardiac • Melanocytes, neurons, cartilage, musculo-connective tissue wall of large arteries • Trunk • Dorsal root ganglia, sympathetic ganglia, adrenal medulla, nerve clusters around aorta • Vagal and Sacral • Parasympathetic ganglia of gut
Trunk Neural Crest Cells: Migration • Dorso-lateral inhibited • At first, between somites, but Semaphorin-3F causes movement through somites.
Further migration • Cells that stop in sclerotome- dorsal root ganglia • Cells past sclerotome-lack wnt and neurotropin receptors- parasympathetic and sympathetic nerves. • BMPs from aorta convert cells to sympathetic and adrenal lineage • Glucocorticoids- adrenomedullary cells. • Neural crest cells committed to path early, but differentiate to determine final location.
Cranial Neural Crest • Head is most anatomically sophisticated portion of vertebrate body. • Mainly composed of neural crest cells
Neural Crest and neurocranium • Viscerocranium (jaws, pharyngeal arches) well established as being of Neural crest origin…neurocranium controversial origin