page_title: Tooth enamel
section_title: Development
hierarchical_section_title: Tooth enamel / Development
caption_attribution_description: English: Histology of developing tooth with enamel organ, dental lamina, and dental papilla labeled.
context_page_description: Tooth enamel is one of the four major tissues that make up the tooth in humans and many other animals, including some species of fish. It makes up the normally visible part of the tooth, covering the crown. The other major tissues are dentin, cementum, and dental pulp. It is a very hard, white to off-white, highly mineralised substance that acts as a barrier to protect the tooth but can become susceptible to degradation, especially by acids from food and drink. In rare circumstances enamel fails to form, leaving the underlying dentin exposed on the surface.
context_section_description: Enamel formation is part of the overall process of tooth development. Under a microscope, different cellular aggregations are identifiable within the tissues of a developing tooth, including structures known as the enamel organ, dental lamina, and dental papilla. The generally recognized stages of tooth development are the bud stage, cap stage, bell stage, and crown, or calcification, stage. Enamel formation is first seen in the crown stage.
  Amelogenesis, or enamel formation, occurs after the first establishment of dentin, via cells known as ameloblasts. Human enamel forms at a rate of around 4 μm per day, beginning at the future location of cusps, around the third or fourth month of pregnancy. As in all human processes, the creation of enamel is complex, but can generally be divided into two stages. The first stage, called the secretory stage, involves proteins and an organic matrix forming a partially mineralized enamel. The second stage, called the maturation stage, completes enamel mineralization.
  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 dentin, allowing for the development of Tomes' processes at the apical pole of the cell. 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 phosphate crystals.
  In the maturation stage, the ameloblasts transport substances used in the formation of enamel. Histologically, 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 final mineralization process compose most of the transported material. The noteworthy proteins involved are amelogenins, ameloblastins, enamelins, and tuftelins. How these proteins are secreted into the enamel structure is still unknown; other proteins, such as the Wnt signaling components BCL9 and Pygopus, have been implicated in this process. During this process, amelogenins and ameloblastins are removed after use, leaving enamelins and tuftelin in the enamel. By the end of this stage, the enamel has completed its mineralization.
  At some point before the tooth erupts into the mouth, but after the maturation stage, the ameloblasts are broken down. Consequently, enamel, unlike many other tissues of the body, has no way to regenerate itself. After destruction of enamel from decay or injury, neither the body nor a dentist can restore the enamel tissue. Enamel can be affected further by non-pathologic processes.
  Enamel is covered by various structures in relation to the development of tooth:
  Nasmyth membrane or enamel cuticle, structure of embryological origin is composed of keratin which gives rise to the enamel organ.
  Acquired pellicle, structure acquired after tooth eruption is composed of food debris, calculus, dental plaque (organic film).
caption: Histologic slide showing a developing tooth. The mouth would be in the area of space at the top of the picture.
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