Dr.

Tahani Abualteen

Development of the root & the periodontium

Root development: Root development starts sometime after enamel completion Enamel formation has to be completed and all the thickness of enamel must be deposited and also fully mature before any part of the root starts to develop For the root to develop there must be an interaction between three components: Dental follicle It is the ectomesenchymal tissue outside the enamel organ and responsible for the formation of the tooth supporting tissues (cementum, PDL and alveolar bone) Epithelial root sheath of Hertwig Epithelial root sheath is an ectodermal structure derived from the cervical loop region of enamel organ (which is a double layered circular structure made of IEE & EEE found at the cervix of the crown after its complete formation) This sheath separates the dental papilla (on the inside) from the dental follicle (on the outside) Dental papilla It is the ectomesenchymal tissue inside the enamel organ and responsible for the formation of dental pulp and dentine ** Both of the crown and root portions of the tooth require an interaction between ectodermal and ectomesenchymal components to form Onset of root development coincides with axial phase of tooth eruption Once the root starts to form and elongate, the tooth starts to erupt (moves vertically up) When the crown starts to have its root, the root elongates (action), and because it can't penetrate the bone underneath it, the tooth erupts (reaction)

This picture is a summary of what we have covered previously: Tooth germ = enamel organ + dental papilla + dental follicle Enamel organ is responsible for the formation of enamel and it is composed of (IEE + EEE + SR + SI)
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Dr. Tahani Abualteen

Tooth germ passes into three stages that are named according to the shape of enamel organ (bud, cap and bell) In the early bell stage morphogenesis 3-dimensional shape of the crown is established In the late bell stage Histogenesis hard tissue formation (enamel and dentin) begins Dentinogenesis starts before amelogenesis When enamel formation is complete, root development begins When root development begins, tooth eruption begins too When tooth completes its eruption and emerges in the mouth, 2/3s of its root is formed, and it needs 2-3 years (if permanent) or 1-1.5 years (if deciduous) to have its root completely formed after emergence into mouth Tooth is only functional, when it makes contact with its opposing tooth (when reaching the occlusal plane) ** Eruption = the full dynamic process from beginning of tooth movements until occlusal contact with opposing tooth is achieved Epithelial root sheath (ERS): ERS forms at the late bell stage from the cervical loop area of enamel organ ERS appears while amelogenesis and dentinogenesis are well-advanced but starts to function in root formation after amelogenesis terminates and completes ERS is a double-layered circular sheath that is composed of: o Internal enamel epithelium (columnar cells from the inside ) o External enamel epithelium (cuboidal cells from the outside ) Unlike enamel organ, the ERS doesn't have stellate reticulum (SR) and stratum intermedium (SI) in between EEE and IEE So in ERS we only have EEE and IEE but we don't have SR and SI and because SR and SI are so important in enamel formation, then NO enamel formation takes place in the root portion of the tooth (enamel is only found in the crown portion of the tooth) So all of the components of enamel organ are important and we should have them all together as a group to be able to form enamel and what confirms this fact is that in the root portion of the tooth we only have EEE and IEE and we don't have SR and SI so that no enamel forms ERS undergoes apical proliferation (ERS proliferates toward the apex of the root and away from the crown) and this is important for mapping out the 3-dimensional shape of the root Coronal proliferation, cell division and movement of IEE cells of enamel organ in the crown is very important for the 3-dimensional structure of the crown Apical proliferation, cell division and movement of IEE cells of ERS in the root is also very important for the 3-dimensional structure of the root
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Dr. Tahani Abualteen

ERS is never a continuous sheath and it undergoes disintegration The ERS cells in the upper part of the root proliferate and form the first part of the root, and as long as this part is being formed, ERS is intact, but once this part is fully formed, the ERS cells disintegrate and disappear and then another ERS cells in a lower part of the root will proliferate and form the second part of the root, and as long as this part is being formed, ERS is intact, but once this part of the root is fully formed, the ERS cells disintegrate and disappear . ERS cells are responsible for the formation of root dentine and once dentine is formed at certain region along the root, these ERS cells disintegrate and the process of root dentine formation continues apically so that each time dentine is formed the ERS cells adjacent to that dentine disintegrate and that's why when we view the root we will NOT see the ERS as a continuous sheath When ERS cells disintegrate, they are removed from the root portion of the tooth & placed into the PDL where they remain as islands of non-functioning cells called (epithelial rests of Malassez) which if proliferate later in response to inflammation, this will give rise to cysts

Enamel pearl: It is a localized area of enamel on root surface Normally we have no enamel formation in the root portion of the tooth because the ERS only has EEE and IEE but NO SR and SI in between

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Dr. Tahani Abualteen

Sometimes the ERS retains the SR and SI in between the EEE and IEE and so a small piece of enamel forms in the root and this piece is called the enamel pearl o Enamel pearls are usually located in the furcation regions of maxillary molars

ERS in multi-rooted teeth: Once enamel formation is completed, ERS starts root development and something called "Primary apical foramen" appears if we view the tooth and the ERS from below If the tooth is single rooted This Primary apical foramen remains as one foramen If the tooth is multi-rooted This primary apical foramen subdivides into a number of "secondary apical foramina" On the right, this is a forming root and a developing ERS as viewed from below: o At first, we will be having a primary apical foramen o If the tooth is single rooted, then this primary apical foramen won't divide and it will proceed as one foramen till apically o If the tooth has two roots , then two horizontal extensions called "epithelial shelves" grow and extend from the margins of ERS toward the center of the root where they fuse and divide the original primary apical foramen into two secondary apical foramina (each of which will continue its root formation on its own) o If the tooth has three roots , then three horizontal extensions called "epithelial shelves" grow and extend from the margins of ERS toward the center of the root where they fuse and divide the original primary apical foramen into three secondary apical foramina (each of which will continue its root formation on its own) o Growth and extension of the epithelial shelves is believed to occur along paths of low vascularity o Number and location of the epithelial shelves: It corresponds to the number of the roots May be under inductive role of dental papilla o If the tooth is multi-rooted and the primary apical foramen has to subdivide into a number of secondary apical foramina then this subdivision doesn't occur at the junction between the crown and the root. We will have a distance between the end of the crown and the beginning of the root division and this distance is called "root trunk" The area where primary apical foramen divides into a number of secondary apical foramina and where a number of epithelial shelves fuse together is called "Furcation area"
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Dr. Tahani Abualteen

Primary apical foramen: While the root is being formed the apex is wide "open" and surrounded by thin regular knife edge of dentine A permanent tooth: o Erupts with about 2/3s of its root formed o Its root needs 2-3 years after eruption to have its root formation completed A deciduous tooth: o Erupts with about 2/3s of its root formed o Its root needs 1-1.5 years after eruption to have its root formation completed By root completion, the wide apical foramen becomes very narrow and only wide to transmit the neurovascular bundle ** How to differentiate between fractured tooth (after root formation completion) and a still developing tooth by looking at its apical foramen?! - Fractured tooth narrow apical foramen - Still developing tooth wide apical foramen Growth of the ERS: Growth and proliferation of ERS occurs apically (toward the apex of the root) ERS is never a continuous sheath so that ERS cells at upper parts of the root form the root and then disintegrate while ERS cells at the lower parts of the root are still forming the root Dental papilla lies internal to ERS and forms dentine and pulp ERS grows apically to enclose the dental papilla except at the apical foramen where the dental papilla is continuous with the dental follicle Margin of ERS is angled internally to form the "Root diaphragm" Dental follicle lies external to ERS and forms cementum, PDL and alveolar bone

Root diaphragm: It is the portion of the ERS that is angled internally toward the dental papilla It is a circular band (with internal & external circles) It is sandwiched between two proliferation zones of undifferentiated mesenchyme: 1. The undifferentiated cells of Dental papilla inside (that will give odontoblasts that will lay down root dentine on the inside) 2. The undifferentiated cells of Dental follicle outside (that will give cementoblasts that will lay down cementum on the outside)
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Dr. Tahani Abualteen

Root diaphragm is the only portion of the ERS that doesn't undergo disintegration and appears always as a continuous band

Commencement of root dentinogenesis & cementogenesis: Commencement = beginning ERS is made up of two layers (internal & external layers which are continuation of IEE & EEE of enamel organ) Cells of internal layer of ERS induce the peripheral undifferentiated cells of the dental papilla to differentiate into odontoblasts to start depositing the root dentine Once the root dentine is laid down, the ERS loses its continuity and disintegrate leading to the formation of the epithelial rests of Malassez in the PDL Disintegration of ERS cells allows the deposited dentine to become exposed to the peripheral undifferentiated cells of the dental follicle which will then differentiate into cementoblasts to start depositing cementum ** Exposure of dentine to dental follicle only occurs in the root but not in the crown ** The process of root dentinogenesis & cementogenesis happens in each piece of the root individually and it isn't neccissary to see them in all areas of the root at once ! o This picture shows the crown and root portions of a developing tooth o Enamel in blow and dentine in brown o Ameloblasts in white and odontoblasts in yellow o Enamel organ = ameloblasts + SI + SR + EEE o Epithelial Root Sheath = EEE and IEE ONLY at the cervical margin of the crown o ERS separates dental papilla on the inside from dental follicle on the outisde cells of internal layer of ERS induce the undifferentiated cells of the dental papilla odontoblasts are formed root dentine is deposited ERS cells adjacent to the deposited dentine disintegrate root dentine becomes exposed to the undifferentiated cells of the dental follicle cementoblasts are formed cementum is deposited
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Dr. Tahani Abualteen

Dental follicle near the diaphragm: Dental follicle near the diaphragm is made of three layers: 1- Inner investing layer: It is the inner layer of the dental follicle which lies in proximity to the epithelial root sheath and the root dentine It is ectomesenchymal in origin (originate from neural crest cells ) Cells in this layer differentiate into cementoblasts (cuboidal cells) which lay down cementum on the top surface of root dentine 2- Intermediate layer: It is the middle layer of the dental follicle It is mesodermal in origin Cells in this layer differentiate into fibroblats which form the periodontal ligament (PDL) PDL is the structure binding the cementum of the root to the alveolar bone 3- Outer layer It is the outer layer of the dental follicle and the farthest away from the epithelial root sheath and the root dentine It is mesodermal in origin Cells in this layer differentiate into osteoblasts which lay down the alveolar bone ** Neural crest cells aren't found in all regions of dental follicle , but they are only found in the inner investing layer ** For deciduous teeth bone is forming as the root is forming BUT for permanent teeth bone is lready formed before root starts to form ** Enamel and epithelial root sheath ectodermal in origin ** Dental pulp and dentine and cementum ectomesenchymal in origin ** PDL and alveolar bone mesodermal in origin Dental Follicle after commencement of cementogenesis: Follicular cells are obliquely (not horizontally) oriented along the root surface Once the root dentine is deposited, epithelial root sheath cells disintegrate & dentine contacts the inner undifferentiated follicular cells which then differentiate into cementoblasts & lay down cementum Intermediate undifferentiated follicular cells differentiate into fibroblasts & start forming PDL o These fibroblasts travel obliquely along the root surface and secrete mostly oblique bundles of collagen fibers which become embedded in the developing cementum from the inside before mineralization and then they get mineralized and they become also embedded in the developing alveolar bone from the outside before mineralization and then they get mineralized o The edges of these bundles which are embedded into cementum from one side and into alveolar bone from the other side and get mineralized are called Sharpys fibers Outer undifferentiated follicular cells differentiate into osteoblasts & start forming alveolar bone

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Dr. Tahani Abualteen

Root completion: In molars of herbivores (e.g. horses, cows, goats and sheep) root formation is a continuous process and apical foramen remains widely open ** Teeth of herbivores are always abraded due to the nature of food they have and this necessitates continuous formation In carnivores & omnivores (e.g. humans) root formation is an ending process and apical foramen needs to narrow sometime ** Teeth of carnivores and omnivores aren't abraded in the same way of herbivores because the nature of food is different and thus they should have some sort of root completion at some time not to end up having occlusal problems (e.g. maxilla and mandible moving away from one another) When the final root length is achieved the proliferation of epithelium in the diaphragm lags behind that of the dental papilla or the dental follicle Final root length is genetically determined inside the ERS itself Root development depends on the interaction between three components (epithelial root sheath that is sandwiched between the dental papilla cells from the inside and the dental follicle cells from the outside) Usually the rate of growth of these three components is the same When the final root length is achieved and root formation is to be terminated, the rate of proliferation of ERS cells slows down so that it becomes slower than the rate of proliferation of dental papilla cells from the inside and dental follicle cells from the outside So that more dentine will be forming from the inside & more cementum will be forming from the outside and this narrows the wide apical foramen & terminates the process of root formation ** Along the length of the root, dentine is always a thin knife edge structure because ERS is moving quickly from one segment of the root to another and forcing odontoblasts to follow it and this doesn't give odontoblasts enough time to build thickness of dentine ** ERS only slows down when final root length is achieved and root formation is to be completed and this gives odontoblasts enough time to build thickness of dentine and narrow apical foramen The wide apical foramen is reduced: o First to the width of the diaphragmatic opening itself o Further narrowed by apposition of dentine and cementum at the apex of the root o Apical foramen should be wide only to transmit the neurovascular bundle

Accessory root canals: They are small secondary canals extending from the main pulpal canal to the surrounding periodontal ligament (PDL) When do these accessory canals occur?! If continuity of epithelial root sheath is broken or not established prior to dentine formation then a defect in the dentinal wall of the pulp occurs Normally, ERS cells disintegrate at certain segment along the root ONLY after complete root dentine formation at that area

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Dr. Tahani Abualteen

If ERS cells disintegrate prematurely or their continuity isn't established (because there was a blood vessel in the area so that they just looped around it) before any root dentine starts to form, then we will have an area that is devoid of dentine and then we will be having an accessory canal extending from the dental pulp (inside) to the PDL (outside) Root elongation & tooth eruption: Let's imagine that my upper body half is the crown and my lower body half is the root then if I want to grow taller, I can't penetrate the ground but instead I can move up The same applies for the tooth, if the root wants to grow longer, then the tooth can't penetrate the bone underneath it but instead it can move upward (trying to erupt) and allowing the root to elongate farther more Notice that root elongation occurs in an opposite direction to tooth eruption Root growth theory of eruption: 1- Old theory is called "cushion hammock ligament" This theory isn't accepted anymore It says that we have a hammock ligament from one edge of the root diaphragm to the other edge This ligament is fibrous with fluid filled interstices and it is attached on either sides to alveolar bone This ligament has a very resistant base thus it prevents bone resorption from below so that the tooth can't penetrate the bone underneath it but instead it will move up (erupts) and allows the root to elongate 2- Current theory is called "pulp limiting membrane" This theory is the one accepted these days This theory rejects and corrects the old theory: They found later that this hammock ligament is not attached to alveolar bone but instead it merges with the developing PDL So that this hammock ligament doesn't have a very resistant base They found that if we remove this hammock ligament surgically, then this will NOT affect the eruption process and the tooth can still move up and erupt So that the hammock ligament is NOT that important in the process of eruption!! The current theory states that there will be changes in vascular permeability in the apical region (between the inside & outside of the tooth) that leads to dense accumulation of tissue fluids that pushes the root and the bone apart and contribute to tooth eruption
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Dr. Tahani Abualteen

Formation of principal PDL fibers: Principal PDL fibers are the first formed fibers in the PDL and they pass into 4 stages

Stage 1 before eruption: o PDL fibers before eruption are loosely structured collagenous elements that aren't making any bundle o In permanent molars: Dentogingival and oblique fibers are organized Dentogingival fibers = fibers connecting the tooth with the gingiva Oblique fibers = fibers extending obliquely from the root surface to the surrounding bone o In premolars: Only dentogingival fibers are well-developed Stage 2 tooth penetration: o Tooth penetration = tooth is moving up & starting to penetrate alveolar mucosa to appear in mouth o PDL fibers during tooth penetration are well differentiated but not spanning the PDL space (they are concentrated cervically but arent found in all parts of the root and aren't connecting the root to the bone yet because the tooth is still erupting and it isnt functioning yet) o In permanent molars: Oblique fibers are well-differentiated but NOT spanning the full thickness of the PDL space o In premolars: Only alveolar crest fibers are organized ** Organization of PDL fibers in molars is more advanced than that in premolars & anteriors Stage 3 reaching occlusion: o Reaching occlusion = the erupting tooth contacts the opposing tooth o In permanent molars: Fibers in the cervical region are organized o In premolars: Fibers in the cervical region are prominent Fibers in the apical part of the root are undeveloped Stage 4 functional: o Functional = erupting tooth reached the opposing tooth and now it is functioning in mastication o It is the only stage where there will be a classical organization of principal fibers of PDL
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Dr. Tahani Abualteen

Additional notes on formation of principal PDL fibers: Collagen fibers are only organized in function (they only gain their classical organization after functioning) Eruptive force caused by traction of collagen in PDL development is debatable One of the theories on tooth eruption and root elongation is that as the PDL tries to develop, the traction of collagen fibers in the PDL leads to eruption This theory is debatable and not 100% clear because collagen fibers in the PDL are only organized after the tooth starts functioning and not before eruption so they can't actually cause the tooth to erupt Change in obliquity & thickness of fibers as the tooth reaches the functional position Obliquity (inclination) of PDL fibers decreases as the tooth reaches its functional position Thickness of PDL fibers increases as the tooth reaches its functional position During eruption resorption of the bone in the base of the socket is predominant If we view a tooth during eruption we will find that bone in the apical region is being resorbed mainly So that bone deposition at the apical region of the tooth is not accepted as a cause of eruption However, species may differ so that (in dogs for example) deposition of bone in the base of the socket is predominant Bone activity is related to the distance the tooth has to erupt Distance greater than root length is associated with bone deposition - Bone resorption at the base of the socket predominates BUT If the distance between the forming tooth and the surface of the oral mucosa is very long and greater than the length of the root of that tooth then we should have deposition of bone at the base of the socket otherwise we will be having a space underneath the root - Like for example the upper canines which have the greatest distance away from the oral mucosa, that's why in such teeth we should have bone deposition near the apex of the tooth to compensate for the upward movement of the tooth - Other teeth are very close to the oral mucosa and thus they don't need bone deposition at the base of the socket but instead they mainly have bone resorption

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