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Transcript
Neurulation and Ectoderm Chapters 12 & 13 Ectodermal Derivatives • • • • • Three main derivatives Neural tube Skin Neural crest Neurulation is process that separates neuralectoderm, epidermal ectoderm, and neural crest I. Neurulation • • • • Formation of Neural Tube Two major processes – – Primary Neurulation: folding of neural plate Secondary Neurulation: cavitation from solid cord Primary Neurulation anterior – – – Fish & frogs: all but tail Birds: anterior to hindlimbs (1-27th somite) Mammals: anterior to sacrum Secondary Neurulation posterior – – – Fish & frogs: tail Birds: posterior to hindlimbs (28th somite on back) Mammals: sacrum and posterior Figure 12.7 Secondary Neurulation in the Caudal Region of a 25-Somite Chick Embryo I . Neurulation • • • • • • • • • Primary Neurulation Shaping & Bending of Neural Plate Closing of Neural Tube Separation from overlying Epidermis Dorso-ventral Patterning Brain Regions Neuronal Development Eye Formation Neuronal Stem Cells 1 I A. Primary Neurulation • • • • Midline cells elongate neural plate – Induction from notochord Lateral cells flatten epidermis Edge of plate folds up (neural folds) Folds move medially & fuse I B. Shaping & Bending of Neural Plate • • • • Median Hinge Point (MHP) – – – Cells anchor to notochord Shorten, become wedge shaped Bends plate inward Dorsolateral Hinge Point (DLHP) – – – – – Cells elongate via microtubules Become wedge shaped (microfilaments at apex) Bends edges inward Colchicine inhibits elongation Cytochalasin B inhibits wedge formation Lateral epidermal ectoderm pushed medially Folds forced together I C. Closing of Neural Tube • • • • Anterior neuropore at front Posterior neuropore at back Failure to close Anterior Neuropore anencephaly Failure to close Posterior neuropore spinal bifida I D. Separation from overlying Epidermis • • • Neural tube cells change CAM’s Prior to neurulation – E-cadherin (Ca++ dependent) During neurulation – – N-cadherin (Ca++ dependent) N-CAM (Ca++ independent) I E. Dorso-ventral Patterning • Dependent upon dual, opposing gradients 2 • • • Ventral floor (& notochord) secretes sonic hedgehog – – Diffuses dorsally, supresses gene expression for dorsal characteristics Causes differentiation of motor neurons Dorsal epidermis cells secrete BMP-4 and BMP-7 – – Counteract sonic hedgehog Stimulates TGF-β paracrine family Adding second notochord induces second set of motor neurons I F. Brain Regions Starts with three primary regions/vesicles • Prosencephalon (forebrain) • Mesencephalon (midbrain) • Rhombencephalon (hindbrain) The 3 regions give rise to 5 secondary vesicles • Prosencephalon (forebrain) • • – – Telencephalon Diencephalon Mesencephalon (midbrain) Rhombencephalon (hindbrain) – – Metencephalon Myelencephalon I G. Neuronal Development Dendrites • Fine outgrowth, receptors During 1st year after birth, enough dendrites form to make 100,000 connections for each cortical neuron • Average cortical neuron connects to 10,000 other neural cells Axons • Long extension of cell body, carry impulse away from cell body • Forms as outgrowth of cell • Elongates along length due to microtubules • – Colchicine causes length regression Microspikes in growth cone area moved by microfilaments – Cytochalasin B blocks microspikes I H. Eye Formation • Outgrowth of diencephalon 3 • • Sonic hedgehog separates eye field into two bilateral fields Mutation of sonic hedgehog can cause cyclopia I I. Neuronal Stem Cells • • • Evidence from mammals and birds that some cells in brain continue to divide after embryonic development BrdU (thymine analog) incorporates in cell undergoing replication (S period) Brains of people, mice, birds treated with BrdU shows uptake and incorporation II. Epidermis • • • • • • Basal layer gives rise to all cells of epidermis Cells keratinize as move to surface Melanin added in lowest two layers 8 weeks to move human cells from basal area to stratum corneum Last 2 weeks in s.c. Note arrow! Two main growth factors stimulate epidermis • TGF-α (transforming growth factor) • – – From basal cells Stimulate own division (autocrine) KGF (keratinocyte growth factor) – – – From underlying dermis (mesoderm) Same as FGF-7 Autocrine or paracrine? III. Neural Crest Cells • • • • Form from edge of neural plate Form from edge of neural plate Migrate laterally and ventrally Cells migrate individually Diverse cell fates • Pigment cells • Nerve cells and ganglia (spinal and autonomic ganglia) • Cartilage and bone, especially of head, neck, face • Adrenal medulla 4 • Connective tissue Cranial Neural Crest • Fates different from trunk neural crest – – Only cranial neural crest form cartilage and bone Contribute to pharyngeal arch tissue Last updated 15 April 2004 5