Amelogenesis

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Amelogenesis is the formation of enamel on teeth and begins when the crown is forming during the advanced bell stage of tooth development after dentinogenesis forms a first layer of dentin. Dentin must be present for enamel to be formed. Ameloblasts must also be present for dentinogenesis to continue.

Contents

A message is sent from the newly differentiated odontoblasts to the inner enamel epithelium (IEE) that causes epithelial cells to further differentiate into active secretory ameloblasts. Dentinogenesis is in turn dependent on signals from the differentiating IEE in order for the process to continue. This prerequisite is an example of the biological concept known as reciprocal induction, in this instance between mesenchymal and epithelial cells.

Stages

Amelogenesis is considered to have three stages. [1] The first stage is known as the inductive stage, the second is the secretory stage, and the third stage is known as the maturation stage. During the inductive stage, ameloblast differentiation from IEE is initiated. Proteins and an organic matrix form a partially mineralized enamel in the secretory stage. The maturation stage completes enamel mineralization.

Inductive (or pre-secretory) stage

In the inductive stage, the morphodifferentiation phase the shape of the crown is determined by the bell stage of tooth development. There is a basal lamina between the IEE and the dental papilla. [2] At this time, the dentin is not mineralized. The IEE cuboidal or low columnar with centralized nuclei and poorly developed Golgi complexes.

The differentiation phase of the Induction stage is initiated by the presence of newly formed predentin. The IEE cells then elongate and become preameloblasts. There is a shift in polarity. Each preameloblast elongates and becomes an postmitotic, polarized, secretory ameloblast. However, there are no Tomes' process yet. It is at this stage that a signal is sent from the newly differentiated ameloblasts back across the dentinoenamel junction (DEJ) to stimulate dentinogenesis.

Secretory stage

In the secretory stage, ameloblasts are polarized columnar cells. In the rough endoplasmic reticulum of these cells, enamel proteins are released into the surrounding area and contribute to what is known as the enamel matrix, which is then partially mineralized by the enzyme alkaline phosphatase. When this first layer is formed, the ameloblasts move away from the interface with dentin, allowing for the development of Tomes' processes at the end of the cell which is in contact with the DEJ. Tomes' process is the term given to the end of the cell which lays down the crystals of the enamel matrix. The Tomes' processes are angled, which introduces differences in crystallite orientation, and hence structure. Enamel formation continues around the adjoining ameloblasts, resulting in a walled area, or pit, that houses a Tomes' process, and also around the end of each Tomes' process, resulting in a deposition of enamel matrix inside of each pit. The matrix within the pit will eventually become an enamel rod, and the walls will eventually become interrod enamel. The only distinguishing factor between the two is the orientation of the calcium crystals.

Maturation stage

In the maturation stage, the ameloblasts transport substances used in the formation of enamel. Microscopically, the most notable aspect of this phase is that these cells become striated, or have a ruffled border. These signs demonstrate that the ameloblasts have changed their function from production, as in the secretory stage, to transportation. Proteins used for the mineralization process compose most of the material transported into the matrix, importantly amelogenins, ameloblastins, enamelins, and tuftelins. The Ca2+ mainly comes from the enamel organ, and not the dental papilla, by either passive, extracellular transportation or active, intracellular transportation. The active route is controlled by ameloblasts so the site of mineralization can have a tightly controlled climate, including modulation of proteins that inhibit mineralization (e.g. Serum-derived Albumin) and concentration of ions.

As enamel is secreted, some mineralisation occurs by Ca2+ deposition between nanospheres of amelogenins forming crystallites. Tuftelin also is suggested to have a role in directing the initial deposition.

The undermineralised, immature enamel containing long, thin prisms of hydroxyapatite, now matures. As the prisms in the enamel grow in thickness but not length, proteins (amelogenins and most non-amelogenins) are removed from the matrix to give more space for hydroxyapatite deposition - mature crystals are hexagonal and 25x75nm and can run the whole length of the enamel (up to 2.5mm). [3] The mineralising enamel becomes progressively less porous. During this process, enamelins and tuftelin are left in the enamel (responsible for enamel tufts).

By the end of this stage, the enamel has completed its mineralization. Enamel mineralization only occurs once (as ameloblasts are lost with eruption within the reduced enamel epithelium); therefore after amelogenesis, enamel production has been finalized. [4] This is in contrast to dentin formation which occurs throughout life (secondary dentin production).

See also

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<span class="mw-page-title-main">Cementum</span> Specialized calcified substance covering the root of a tooth

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<span class="mw-page-title-main">Dentin</span> Calcified tissue of the body; one of the four major components of teeth

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Enamelin is an enamel matrix protein (EMPs), that in humans is encoded by the ENAM gene. It is part of the non-amelogenins, which comprise 10% of the total enamel matrix proteins. It is one of the key proteins thought to be involved in amelogenesis. The formation of enamel's intricate architecture is thought to be rigorously controlled in ameloblasts through interactions of various organic matrix protein molecules that include: enamelin, amelogenin, ameloblastin, tuftelin, dentine sialophosphoprotein, and a variety of enzymes. Enamelin is the largest protein (~168kDa) in the enamel matrix of developing teeth and is the least abundant of total enamel matrix proteins. It is present predominantly at the growing enamel surface.

<span class="mw-page-title-main">Ameloblastin</span> Protein-coding gene in the species Homo sapiens

Ameloblastin is an enamel matrix protein that in humans is encoded by the AMBN gene.

<span class="mw-page-title-main">Tuftelin</span> Protein-coding gene in the species Homo sapiens

Tuftelin is an acidic phosphorylated glycoprotein found in tooth enamel. In humans, the Tuftelin protein is encoded by the TUFT1 gene. It is an acidic protein that is thought to play a role in dental enamel mineralization and is implicated in caries susceptibility. It is also thought to be involved with adaptation to hypoxia, mesenchymal stem cell function, and neurotrophin nerve growth factor mediated neuronal differentiation.

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<span class="mw-page-title-main">Dental papilla</span>

In embryology and prenatal development, the dental papilla is a condensation of ectomesenchymal cells called odontoblasts, seen in histologic sections of a developing tooth. It lies below a cellular aggregation known as the enamel organ. The dental papilla appears after 8–10 weeks intra uteral life. The dental papilla gives rise to the dentin and pulp of a tooth.

<span class="mw-page-title-main">Odontoblast</span> Type of cell that produces dentin in teeth

In vertebrates, an odontoblast is a cell of neural crest origin that is part of the outer surface of the dental pulp, and whose biological function is dentinogenesis, which is the formation of dentin, the substance beneath the tooth enamel on the crown and the cementum on the root.

Dentinogenesis is the formation of dentin, a substance that forms the majority of teeth. Dentinogenesis is performed by odontoblasts, which are a special type of biological cell on the outer wall of dental pulps, and it begins at the late bell stage of a tooth development. The different stages of dentin formation after differentiation of the cell result in different types of dentin: mantle dentin, primary dentin, secondary dentin, and tertiary dentin.

An odontoblast process is an extension of a cell called an odontoblast, which forms dentin in a tooth. The odontoblast process is located in dentinal tubules. It forms during dentinogenesis and results from a part of the odontoblast staying in its location as the main body of the odontoblast moves toward the center of the tooth's pulp. The odontoblast process causes the secretion of hydroxyapatite crystals and mineralization of the matrix secreted by the odontoblasts.

<span class="mw-page-title-main">Enamel tufts</span>

Enamel tufts are hypomineralized ribbon-like structures that run longitudinally to the tooth axis and extend from the dentinoenamel junction (DEJ) one fifth to a third into the enamel. They are called ‘‘tufts’’ due to their wavy look within the enamel microstructure.

Dentin sialophosphoprotein is a precursor protein for other proteins found in the teeth. It is produced by cells (odontoblasts) inside the teeth, and in smaller quantities by bone tissues. It is required for normal hardening (mineralisation) of teeth. During teeth development, it is broken down into three proteins such as dentin sialoprotein (DSP), dentin glycoprotein (DGP), and dentin phosphoprotein (DPP). These proteins become the major non-collagenous components of teeth. Their distribution in the collagen matrix of the forming dentin suggests these proteins play an important role in the regulation of mineral deposition. Additional evidence for this correlation is phenotypically manifested in patients with mutant forms of dentin sialophosphoprotein. Such patients suffer dental anomalies including type III dentinogenesis imperfecta.

<span class="mw-page-title-main">Amelogenesis imperfecta</span> Genetic disorder resulting in abnormal enamel

Amelogenesis imperfecta (AI) is a congenital disorder which presents with a rare abnormal formation of the enamel or external layer of the crown of teeth, unrelated to any systemic or generalized conditions. Enamel is composed mostly of mineral, that is formed and regulated by the proteins in it. Amelogenesis imperfecta is due to the malfunction of the proteins in the enamel as a result of abnormal enamel formation via amelogenesis.

Hard tissue, refers to "normal" calcified tissue, is the tissue which is mineralized and has a firm intercellular matrix. The hard tissues of humans are bone, tooth enamel, dentin, and cementum. The term is in contrast to soft tissue.

References

  1. Nanci, Antonio (2013). Ten Cate's Oral Histology. London: Elsevier. pp. 133–148.
  2. Bath-Balogh, Mary; Fehrenbach, Margaret J. (2011). Illustrated Dental Embryology, Histology, and Anatomy. St. Louis, MO: Elsevier. p. 58.
  3. Ten Cate's Oral Histology, Nanci, Elsevier, 2013, page 149-154
  4. Max A. Listgarten, University of Pennsylvania and Temple University, http://www.dental.pitt.edu/informatics/periohistology/en/gu0302.htm