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Amelogenesis 1

The cells forming enamel, the ameloblasts differentiate from the inner enamel epithelium. Ameloblasts begin enamel deposition after a few micrometers of dentin have been deposited. Dentinogenesis begins at the late bell stage of tooth development. Shortly thereafter, amelogenesis begins. The life cycle of the ameloblasts In the early development, the inner dental epithelium is formed of single layer of cuboidal cells supported by a basement membrane which separates them from the dental papilla. These cells undergo differentiation to become active secretory ameloblasts. They, thus exhibit a number of morphologic changes as they differentiate and pass through the following functional stages. 1 !orphogenic " #rgani$ing % Secretory & !aturative ' (rotective ) Desmolyfic

1 !orphogenic stage *

The cells of the inner dental epithelium interact with the ad+acent mesenchymal cells determining the shape of the dentino enamel +unction of the crown.

The ad+acent pulpal layer is a cell free $one contains fine argyrophilic fibers and cytoplasmic processes of the superficial cells of the dental papilla.

During this stage, the cells assume columnar shape with large oval nuclei that almost fill the cell body. ,olgi apparatus and the centerioles are located in the proximal end of the cell -facing the stratum intermedium. while the mitochondria are dispersed throughout the cytoplasm.

/ 0ife cycle of amelobiasts.

The cells become tall columnar and the cell free $one between it and the dental papilla disappears.

Thus the epithelial cells become in close contact with the connective tissue cells of the dental papilla which differentiate into odontoblasts. The odontoblasts begin to secrete dentin during the terminal phase of the organi$ing stage.

A reversal of the functional polarity of these cells ta1e place by the migration of the centriols and ,olgi regions from the proximal ends of the cells into their distal ends. The nucleus free $one at the distal end of the cell become almost as long as the proximal parts containing the nucleus and mitochondria.

There are desmosomal attachment at the proximal and distal part of the cells.

The Secretory stage

The ameloblasts enter their secretory stage after the formation of a layer of dentin.

The ameloblast becomes a highly polari$ed cell with the ma+ority of its organelles situated in the cell body distal to the nucleus. The ,olgi complex increases in volume and the amount of rough endoplasmic reticulum and cytoplasmic inclusions are significantly increased.

The newly formed ameloblasts begin to secrete enamel matrix which constitute the non prismatic layer near the D23.

As the first increment of enamel is formed, the ameloblasts migrate away from the dentin surface and each ameloblast develops a short conical pro+ection 1nown as Tomes / process. A distinction between the process and cell body is mar1ed by the distal terminal bar. The process only contains secretory granules, while the ameloblast cytoplasm contains abundant synthetic organelles.

Tomes /processes extend into the newly forming enamel giving the +unction between ameloblasts and enamel a Saw tooth li1e appearance.

At the end of the enamel matrix formation there is a brief transitional phase involving a reduction in height of the amelobiasts and a decrease in their organelle4s content.

Tomes /processes at the distal ends of the amelobiasts disappear.

During this period most probably the outer structureless or prismless enamel is formed. secretory amelobiasts showing five substages. A,, absorption granules5 A(, apical contact speciali$ation -hemidesosomes.5 Av, autophgagic vacuales #ysosomss.5 6T3, butb type of contacts5 78, coaled -absorplive9. vesicles5 D, desmosomes5 D,, dense -secretory. granules5 ,, ,olgi apparatus5 ,2:, granular -rough. endoplasmic reticulum5 ,r, pale -secretory9. granules5 01.0", 0%, lysosomes5 0,, lipid granules5 !, mitochondria5 !,, mitochondrial granules5 S6, striated border5 T6, terminal bars5 T3, tight +unctions5 T;. terminal web.

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!aturative stage

As the amelobiasts complete the matrix deposition phase, they undergo significant ultrastructural changes lin1ed to the assumption of their new function of enamel maturation.

2xcess organelles associated with synthesis are enclosed in autophagic vacuoles and digested by lysosomal en$ymes.

This development is followed by a shift in many of the remaining organelles to the distal part of the cell and a complex folding of the distal plasma membrane to form a ruffle border.

This ruffle border greatly increases the surface area of the extremity of the ameloblast and indicates that rapid transport of material is ta1ing place across the plasma membrane. 7ytoplasmic vacuoles containing material resembling enamel matrix are present in the cell body. As the enamel maturation nears completion, the amelobiasts lose their ruffle borders and secrete a material between the now flattened distal end of the cell and the enamel surface. This material appears morphologically identical to a basal lamina. <emidesmosomes also form along the distal cell membrane at this time.

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Trotective stage

Then the amelobiasts loose their regular appearance and become cuboidal and often can no longer be differentiated from the cells of the other layers of the epithelial dental organ. These layers together form a stratified epithelial covering of the enamel, the so called :educed Denial 2pithelium -:educed 2namel 2pithelium.. The reduced dental epithelium protects the enamel from the surrounding connective tissue until tooth eruption. If the connective tissue comes in contact with the enamel, enamel can be either resorbed or covered by a layer of cementum.

) Desmolytic Stage* At the time of eruption, the reduced dental epithelium proliferates and seem to elaborate en$ymes -desmolytic en$ymes. that induce atrophy and destruction of the connective tissue separating it from the oral epithelium, so fusion of the two epithelia can occur to form the attachment epithelium. (remature degeneration of the reduced dental epithelium may also prevent tooth eruption.