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Significance of Neural Crest in Tooth Development

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Significance of Neural Crest in Tooth Development OMPJ VP Jayasekharan et al 10.5005/jp-journals-10037-1018 REVIEW ARTICLE Signifi cance of Neural Crest in Tooth Development: The Molecular Signature 1VP Jayasekharan, 2Jacob Kurien, 3Eapen Cherian, 4Renji K Paul, 5Aravind S Raju ABSTRACT cells to those of mesenchymal cells. In the head region, The neural crest originates from cells located along the lateral incipient neural crest cells send out processes that penetrate margins of the neural plate. Neural crest cells arise as the result the basal lamina underlying the neuroepithelium just before of an inductive action by the non-neural ectoderm adjacent to neural tube closure. the neural plate and possibly by nearby mesoderm as well. As the neural tube forms, a group of cells separate from the neuro- The neural crest provides cells for the development ectoderm. These cells have the capacity to migrate and differen- of future organs and tissues. After the induction of neural tiate extensively within the developing embryo and they are the crest, the neural crest cells are formed within neural plate basis of structures such as spinal sensory ganglia, sympathetic border regions, which rise as neural folds, and ultimately neurons, Schwann cells, pigment cells and meninges. Specifi c interactions occur during the development of tooth and recent converge to form the dorsal midline of the neural tube, and research has concentrated more on the molecular aspects of it is from here that the neural crest cells will emerge in an these interactions. Thus, it is highly imperative to understand and anterior to posterior sequence. During and after migration, digress the complex mechanisms involved in these processes. cells differentiate into a wide range of derivatives that are Keywords: Epithelial-Mesenchymal transition, Neuroectoderm, grouped into: (A) ectomesenchymal category and (B) non- Placode. ectomesenchymal category. How to cite this article: Jayasekharan VP, Kurien J, Cherian E, Paul In the last stage, the mammalian teeth develop from two RK, Raju AS. Signifi cance of Neural Crest in Tooth Development: The Molecular Signature. Oral Maxillofac Pathol J 2014;5(2): type of cells: stomodeal ectoderm, which form ameloblasts 484-487. and cranial neural crest-derived (ecto) mesenchymal cells, 2 Source of support: Nil which fi nally form odontoblasts and cementoblasts. Confl ict of interest: None BRANCHIAL ARCH FORMATION INTRODUCTION The accumulation and proliferation of cranial neural crest cells, together with proliferation of the ectoderm cells, The neural crest cells are a unique population of cells that produce a series of swellings that constitute the facial pri- arise from the dorsal part of the forming neural tube during mordia. Mammalian teeth develop on two of these primordia: neurulation. These cells undergo an epithelial to mesen- 1. Frontonasal process. chymal transformation as they detach from the neural tube. 2. First branchial (pharyngeal) arch which forms mandible Neural crest cells differentiate fi rst in the mesencephalic zone and proximal maxilla. of the future brain. In the spinal cord portion of the neural These primordial structures form as cranial neural crest tube, the neural crest cells at the end detach after the neural cells from the developing brain and accumulate beneath folds have fused. Neural crest cells at the very caudal end ectoderm, covering what will become the face and oral of the neural tube are formed from the medullary cord after cavity. In addition, more caudal swellings are also created 1 the caudal neuropore closes on. The neural crest cells bread which form the 2nd, 3rd, 4th branchial arches, each of which free from the neural plate or neural tube by changing their develop into a defi ned set of structures.3 shape and properties from those of typical neuroepithelial SCENARIO FOR THE ORIGIN OF 1,4,5 2 3 Senior Lecturer, Professor and Head, Reader NEURAL CREST 1-3Department of Oral Pathology and Microbiology, St Gregorios Dental College, Kothamangalam, Kerala, India Neural crest evolved from Rohon-Beard cells, a class of primary sensory neurons that occur in dorsal spinal cord of 4,5Department of Orthodontics, St Gregorios Dental College Kothamangalam, Kerala, India chordates, projecting axons peripherally onto lateral migra- Corresponding Author: VP Jayasekharan, Senior Lecturer tion pathway between somites and epidermis that is also Department of Oral Pathology and Microbiology, St Gregorios used by vertebrate neural crest cells. Under the developing Dental College, Kothamangalam, Kerala, India, Phone: scenario, peripheral Rohon-Beard cells would need to dediffe- 8547055944, e-mail: [email protected] rentiate and divide to form sensory neurons and glia of 484 OMPJ Signifi cance of Neural Crest in Tooth Development: The Molecular Signature the dorsal root ganglia and neurons of the central nervous The special sensory nerves and associated glia and ganglia system are not known to dedifferentiate. also arise from placodes. Cranial nerve I arises from the The evolutionary developmental origin of neural crest, olfactory placodes, cranial nerve II arises from optic cup specifi cally, the sensory neurons and glia of dorsal root (distal end of which thickens as the placode-like rudiment ganglia, is to be found in migration of mitotically active of neural retina) and cranial nerve VIII (vestibulococh- Rohon-Beard progenitor cells from dorsal neural tube. Neural lear nerve) and vestibulocochlear ganglion arises from the crest cells would be considered a subset of Rohon-Beard otic placode.8 precursors that delaminated form nascent spinal cord and Neural crest is inhibited from forming in the anterior migrate into the periphery.4 neural ridge by the signaling molecule Dickkopf 1, a Wnt inhibitor that is secreted by prechordal mesoderm. Neural THE MIGRATION OF NEURAL CREST crest cells emerging from the diencephalon posteriorly through r2 do not express any Homeobox gene, (Hox gene). After leaving the neuroepithelium, the neural crest cells fi rst These genes are characterized by possession of a particular encounter a relatively cell-free environment rich in extra- DNA sequence, the homeobox, encoding a variable protein cellular matrix molecules. In this environment, they undergo domain called as homeodain, whereas the cells emerging extensive migrations along several well-defi ned pathways. from the hindbrain region from r3 and posteriorly express These migrations are determined by intrinsic properties of a well-ordered sequence of Hox genes. the neural crest cells and features of the external environ- It is known that some of the neural crest cells associated ment encountered by the migrating cells.5 The migration of with r3 and r5 undergo apoptosis because of the presence neuroectodermal cells lying along the crest of each neural of the apoptosis-inducing molecule BMP-4. A close relation fold occurs ventrolaterally on each side of the neural tube, exists between the pattern of migration of rhombomeric forming an irregular elongated mass between the neural tube neural crest cells and the expression of products of the and overlying surface ectoderm. In addition, cranial and more Hox-b gene complex. Hoxb-2, Hoxb-3, Hoxb-4 products caudal neural crest cells give rise to some identical cell types are expressed in a regular sequence in the neural tube and (neurons) but also some different cell types.6 the neural crest-derived mesenchyme of the second, third Before neural crest emigrates from the neural tube, they and fourth pharyngeal arches. Isthmic tissue emits a high undergo a process called epithelial mesenchymal transition concentration of FGF-8 which suppress Hoxa-2, the selector (EMT) that will confer to them the ability to migrate. A gene for determining the character of the second arch. The similar EMT process in different cellular machineries also introduction of FGF-8 alone into region of future second arch occurs. When the EMT is complete, the neural crest cells suppresses its formation, and it takes on the character of fi rst delaminate from neuroepithelia and migrate along specifi c arch. The cranial mesoderm is presently suspected to play a pathways.7 formerly unexpected role in regulating the fate of migrating neural crest cells. Cranial neural crest cells differentiate into CRANIAL NEURAL CREST CELLS a variety of tissues, including connective tissue and skeletal Neural crest cells from the caudal prosencephalon (forebrain) tissues. These tissues constitute much of the soft and hard and mesencephalon (midbrain) regions give rise to the tissues of the face.5 parasympathetic ganglion of cranial nerve III, a portion of the connective tissue around the developing eyes and optic THE NEURAL CREST—MOLECULAR nerves, the muscles of the iris and ciliary body, and part of LEVEL SIGNATURE the cornea of the eye, they also contribute, along with head Expression of neural crest cell marker genes are preceded mesoderm, to the head mesenchyme cranial to the level of by expression of other genes important in neural crest cell mesencephalon. Neural crest cells from the mesencephalon induction which are expressed in neural plate, msx1 and and rhombencephalon (hindbrain) regions also give rise to msx2 which are expressed in neural folds and non-neural structures in the developing pharyngeal arches of the head ectoderm and Dlx, which is expressed in non-neural ectoderm and neck. The rhombencephalic neural crest cells also contri- only. Together these factors induce expression of a number of bute to some of the cranial nerve ganglia. Specifi cally, rhom- genes and transcription factors including Ap2, cMyc, Foxd3, bencephalic neural crest cells give rise to some neurons Id, Twist and Sox 9 and Sox 10 in a subset of neural plate and all glial cells in the sensory ganglia of cranial nerves V, border cells to form premigratory neural crest cells (Table 1). VII, IX and X. The remaining neurons in sensory ganglia of Both gene and gene-network surveys for neural crest cranial nerves V, VII, IX and X arise from small ectodermal have shown Bmp2 and 4 expressed in non-neural ectoderm, placodes, called epibranchial or epipharyngeal placodes.
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