Amelogenesis is the formation of enamel on

teeth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, t ...

and begins when the crown is forming during the advanced bell stage of

tooth development Tooth development or odontogenesis is the complex process by which teeth form from embryonic cells, grow, and erupt into the mouth. For human teeth to have a healthy oral environment, all parts of the tooth must develop during appropriate st ...

after

dentinogenesis {{Refimprove, date=September 2014 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, ...

forms a first layer of dentin. Dentin must be present for enamel to be formed.

Ameloblasts Ameloblasts are cells present only during tooth development that deposit tooth enamel, which is the hard outermost layer of the tooth forming the surface of the crown. Structure Each ameloblast is a columnar cell approximately 4 micrometers in d ...

must also be present for dentinogenesis to continue. A message is sent from the newly differentiated odontoblasts to the

inner enamel epithelium The inner enamel epithelium, also known as the internal enamel epithelium, is a layer of columnar cells located on the rim nearest the dental papilla of the enamel organ in a developing tooth. This layer is first seen during the cap stage, in wh ...

(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 Mesenchyme () is a type of loosely organized animal embryonic connective tissue of undifferentiated cells that give rise to most tissues, such as skin, blood or bone. The interactions between mesenchyme and epithelium help to form nearly every ...

and epithelial cells.

Stages

Amelogenesis is considered to have three stages. 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. 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 Ameloblasts are cells present only during tooth development that deposit tooth enamel, which is the hard outermost layer of the tooth forming the surface of the crown. Structure Each ameloblast is a columnar cell approximately 4 micrometers in ...

. 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 The endoplasmic reticulum (ER) is, in essence, the transportation system of the eukaryotic cell, and has many other important functions such as protein folding. It is a type of organelle made up of two subunits – 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

amelogenin Amelogenins are a group of protein isoforms produced by alternative splicing or proteolysis from the ''AMELX'' gene, on the X chromosome, and also the ''AMELY'' gene in males, on the Y chromosome. They are involved in amelogenesis, the developme ...

ameloblastin Ameloblastin (abbreviated AMBN and also known as Sheathlin or Amelin) is an enamel matrix protein that in humans is encoded by the AMBN gene. Function Ameloblastin is a specific protein found in tooth enamel. Although less than 5% of enamel con ...

enamelin 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 ame ...

s, and

tuftelin 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 ...

s. The Ca2+ mainly comes from the

enamel organ The enamel organ, also known as the dental organ, is a cellular aggregation seen in a developing tooth and it lies above the dental papilla. The enamel organ which is differentiated from the primitive oral epithelium lining the stomodeum.The enam ...

, and not the

dental papilla 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 ...

, 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). 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.Max A. Listgarten, University of Pennsylvania and Temple University, http://www.dental.pitt.edu/informatics/periohistology/en/gu0302.htm This is in contrast to dentin formation which occurs throughout life (secondary dentin production).

References

Tooth development

Stages

Inductive (or pre-secretory) stage

Secretory stage

Maturation stage

See also

References