FERTILIZATION The human somatic (body) cell contains 46 chromosomes, 46 being the diploid numbers for the cell. Two of these are sex chromosomes ; the remainder are autosomes Each chromosome is paired so that every cell has 22 homologous set of paired autosomes,with one chromosome derived from mother & one from father. Fertilization is the fusion of male & female germ cells to form zygote. The process that produces male & female germ cells(the spermatozoa & ova,collectively called gametes) with half the number of chromosome of the somatic cell is called meiosis. Prenatal development
Divided into three successive phases.
The first two constitute the embryonic stage,& the third is fetal stage. The first phase begins at fertilization & spans the first 4 weeks or so of development.It is mostly characterized by cellular proliferation & migration,with some differentiation. The second phase spans the next 4 weeks of development & is characterized largely by differentiation of all major external & internal structures. From the end of second phase to term,further development is largely a matter of growth & maturation,& the embryo is now called a fetus. Formation of three layered embryo
After fertilization ,mamalian development involves a
phase of rapid proliferation & migration of cell with little or no differentiation. This proliferative stage last untill three germ layers have been formed. So, fertilized egg initially undergoes a series of rapid division that lead to formation of ball of cell,called as morula. Later it transforms into fluid filled hollow ball,the blastocyst. Two cell population can be recognized : those lining the cavity(primary yolk sac),called trophoblast cells,& small cluster within the cavity called the embryoblast. Embryoblast forms embryo proper 8th day of gestation: embryoblast differentiate into two layered disk ,called bilaminar germ disk. The cells situated dorsally,or ectodermal layer ,are columanr & reorganize to form the amniotic cavity. Those on the ventral aspect,the endodemal layer,are cuboidal & form the roof of a second cavity(the secondary yolk sac) Later the axis of the embryo is established& is represented by a slight enlargement of ectodermal & endodermal cells at the head(or rostral) end of the embryo in a region called as prochordal plate. During the third week of development the bilaminar embryonic disk is converted into trilaminar disk. Floor of the amniotic cavity is formed by the ectodermal layer of the bilaminar embryo. The primitive streak is a narrow groove terminating in a circular depression called the primitive node. Surface ectodermal cells migrate to the streak between the ectodermal and endodermal layers to form a solid column.. Through canalization of this cord of cells,the notochord is formed. Space between the newly embryonic endoderm & ectoderm form,called as mesoderm. The cells that accumulate anterior to the prochordal plate as a result of migration of ectodermal cells gives rise to cardiac plate.,the structure in which heart forms. Formation of neural tube & the fate of germ cells
Events during the first three weeks of development
leads to formation of triple layered.or triploblastic embryo. During the next 3 – 4 weeks of development,major tissues & organs differentiates from the triploblastic embryo; these included the head & face & the tissues contributing to the development of teeth. The nervous system develops as a thickening with in the ectodermal layer at the rostral end of the embryo. The thickening constitute the neural plate. Folding of the embryo
Folding of embryo occurs in two planes i.e
rostrocaudal & lateral axis. The head fold is critical for the formation of a primitive stomatodeum or oral cavity; through this fold ectoderm come to line stomatodeum,with the stomatodeum separated from the gut by buccopharyngeal membrane. The Neural Crest
As the neural tube forms,a group of cells separate
from the neuroectoderm. These cells have the capacity to migrate & differentiate extensively within the developing embryo & they are basis for such structures as the spinal sensory ganglia,sympathetic neurons etc. In the avian emryo these cells can be distinguished differentiating & separating at the crest of the neural folds.& hence the name. Neural crest cell in the head region not only helps in formation of cranial sensory ganglia,they also differentiate to form most of the connective tissue of the head. Embryonic connective tissue elsewhere is derived from mesoderm & is known as mesenchyme,whereas in trhe head it is called as ectomesenchyme reflecting its origin from neuroectoderm All the tissues of the tooth (except enamel) & its supporting structures are derived directly from neural crest cells. DEVELOPMENT AND GROWTH OF TEETH Primitive oral cavity or stomodeum is lined by stratified squamous epithelium called as oral ectoderm. At the junction of ectoderm endoderm of foregut is ‘buccopharyngeal membrane’. About 27 day of I.U life it rupturs. Connective tissue cells underlying ectoderm are neural crest or ectomesenchyme in origin. These cell induce the overlying ectoderm to start tooth development. Primary Epithelial Band
Two or three weeks after the rupture of the
buccopharyngeal membrane,when the embryo is 6 week old,certain ares of basal cells proliferate & a continuous band of thickened epithelium forms around the mouth in the presumptive upper & lower jaws. These bands are roughly horseshoe shaped & correspond in position to the future dental arches of the upper & lower jaws Dental lamina
2-3 weeks after rupture of buccopharyngeal
membrane , when embryo is 6 weeeks old,certain areas of basal cells of the oral ectoderm proliferate more rapidly than do the cells of the adjacent part. This leads to formation of primary epithelial band which is a band of epithelium that has invaded into underlying ectomesenchyme along each of the horse –shoe shaped future dental arches. The formation of these bands is the result not so much of increased proliferative activity within the epithelium as of a change in the orientation of the mitotic spindle & cleavage plane of dividing cells. At about seventh week the primary epithelial band divides into an lingual process called as dental lamina & buccal (outer) process called as vestibular lamina Sagittal section through the head of an embryo. A, The thickened epithelium of the primary epithelial band. B, The same structure at higher magnification. C, The change in plane of cleavage. Dental lamina serves as a primordium for the ectodermal portion of the deciduous teeth. Distal extention give rise to permanent molar Lingual extention / successional lamina give rise to rest other permanent teeth. Activity last over a period of at least 5 years. As the tooth continue to develop ,they loose connection with dental lamina. They later break up by mesenchymal invasion. Remnants of dental lamina persist as epithelial pearls or islands with in the jaw or in the gingiva,called as ‘cell rest of serres’ Vestibular lamina
Develop labial to the dental lamina in each dental
arch as epithelial thickening. Also called as ‘Lip furrow band’. Later it hollows & forms oral vestibule between alveolar portion of the jaws & the lips & cheeks. Tooth Development
At certain points along the dental lamina ,the
ectodermal cells multiply still more rapidly & form little knobs that grow into underlying mesenchyme. These knobs represent the beginning of the ‘enamel organ’ of the tooth bud of a deciduous tooth. Development of tooth results from interaction of the epithelium derived from the first arch & ectomesenchymal cells derived from neural crest cells Developmental Stages
Odontogenesis is devided into several morphological
stages ,named after shape of the enamel organ. Bud Stage
In the bud stage enamel organ consist of peripherally
located low columnar cells & centrally located polygonal cells. Many cells of tooth bud & surroundin ectomesencyme undergo mitosis. As a result of that the ectomesenchymal surrounding the tooth bud condenses. This condensed ectomesenchymal cell immediately subjecent to tooth bud is called as dental papilla. The condensed ectomesenchyme that surroun the tooth bud & dental papilla is called as dental sac. Both become well defined as cap & bell stage is reached. Cap Stage
Unequal growth in different parts of tooth bud leads
to cap stage,which is characterized by shallow invagination on the deep surface of the bud. Outer enamel epithelium:Peripheral cells of the cap stage are cuboidal,cover the convexity of the “cap”.Separated from dental sac by delicate basement membrane. Inner enamel epithelium: These are cell present in the concavity of the “cap” become tall,columnar. Stellate reticulum:Polygonal cells present in the centre begin to separate due to water being drawn into the enamel organ from the surrounding dental papilla as a result of osmotic force exerted by glycosaminoglycans contained in the ground substance. As a result the polygonal cell become star shaped but maintain contact with each other by their cytoplasmic process. As these star shaped cells form a cellular network,they are called the Stellate reticulum. Acts as a shock absorber & support & protect the delicate enamel forming cells. Dental papilla:Under the organizing influence of the proliferating epithelium of the enamel organ,the ectomesenchyme that is partially enclosed by the invaginated portion of the inner enamel epithelium proliferates.It condense to form dental papilla. Formative organ of dentin & primordium of pulp. Dental sac(Dental follicle):These result due to marginal condensation of ectomesenchyme,surrounding enamel organ & dental papilla. Bell Stage
As the invagination of epithelium deepens & its
margin continue to grow ,the enamel organ assumes a bell shape. Crown shape is determined . The folding of enamel organ to cause different crown shape is shown to be due to differential rates of mitosis and differences in cell differentiation time The inner enamel epithelial cells which lie in the future cusp tip or the incisal region stop dividing earlier & begin to differentiate first. The pressure exerted by continuous cell division on these differentiating cells from other areas of the enamel organ causes the cell to be pushed out into the enamel organ in the form of cusp tip. Cell proliferation & differentiation occur gradually from cusp tip to the depth of sulcus. The determination of crown shape is under the control of genes& their signalling molecule & growth factor. IEE:Consist of single layer of cells that differentiate prior to amelogenesis into tall columnar cells called ameloblast.4-5µm in diameter & 40 µm in height. OEE:Flatten to low cuboidal form. Dental lamina:Seem to extend lingually & is termed successional dental lamina. Dental papilla:Before the IEE begins to produce enamel,the peripheral cells of mesenchymal papilla differentiate into odontoblast. Stratum intermedium: few layer of squamous cell. Lie between IEE & stellate reticulum. The well developed cytoplasmic organelles ,acid mucopolysaccharides & glycogen deposit indicates a high degree of metabolic activity. This layer seem to be essential for enamel formation. Stellate Reticulum:Expands further. Before enamel formation begins it collapses ,reducing the distance between centrally situated ameloblast & nutrient capillaries near OEE.
The basement membrane that separates the enamel
organ & the dental papila just prior to dentin formation is called the membrana preformitiva. Advanced Bell stage
Characterized by commencement of mineralization &
root formation. The boundary between IEE & odontoblast outlines the future dentinoenamel junction. The formation of dentin occurs first as a layer along the future DEJ in the region of future cusp & proceeds pulpally & apically. After the first layer of dentin is already formed ,the ameloblast which has already differentiated from IEE lay down enamel over the dentin in the future incisal & cuspal areas. The cervical portion of enamel organ give rise to hertwig epitheial root sheaths(HERS). Transitory Structure
1. Enamel niche: The enamel organ may be seen to
have a double attachment of dental lamina to the overlying oral epithelium enclosing ectomesenchyme called enamel niche between them.This appearance is due to funnel shaped depression of the dental lamina. 2. Enamel Knot: The cells in the centre of the enamel organ are densely packed are called as Enamel knot. 3.Enamel cord: vertical extension of enamel knot. 4.Enamel septum: When the enamel cord extends to meet the outer enamel epithelium. 5. Enamel navel: The outer enamel epithelium at the point of meeting shows a small depression & is called as enamel naval. HERS & Root Formation
Development of the root begins after enamel &
dentin formation has reached the future cemento enamel junction. HERS consist of OEE & IEE only.The inner layer cells normally do not produce enamel & remain short. When these cells have induced the differentiaion of radicular dental papilla into odontoblast & first layer of dentin has been laid down,the HERS loses its structural continuity. Photomicrographs summarizing root formation. A, The root is beginning to form as an extension of the inner and outer dental epithelia in the cervical loop region (arrowhead), which form a bilayered structure called Hertwig’s epithelial root sheath. The root sheath will induce differentiation of odontoblasts from the radicular pulp. B, The differentiation of odontoblasts and the formation of root dentin are shown. C, Root formation is almost complete. The root sheath is now a small tag of epithelium that is inducing the final differentiation of odontoblasts at the rim of the future apical foramen Its remenants remnants persist as an epithelial network of strands or clumps near the external surface of the root. These epithelial remanants are found in peridontal ligament of erpted teeth & are called rest of malassez. The differentiation of odontoblast & formation of dentin follow the lengthening of the root sheath. At the same time cells of dental sac proliferates & invades the epithelial layer. The epithelium is moved away from the surface of dentin so that connective tissue cells come into contact with outer surface of the dentin & differentiate into cementoblast that deposit a layer of cementum onto surface of dentin. The outer & inner enamel epithelia bend at the future CEJ into a horizontal plane ,narrowing the wide cervical opening of the tooth germ. Differential growth of the epithelial diaphragm in multi rooted teeth causes the division of the root trunk into two or three roots. During the general growth of the enamel organ the expansion of its cervical opening occur in such a way that long tongue like extension of the horizontal diaphragm develop. If cells of the HERS remain adherent to the dentin surface ,they may differentiate into fully functioning ameloblast & produce enamel. Such droplets of enamel are called as enamel pearl. HISTOPHYSIOLOGY A number of physiologic growth process participate in the progressive development of the teeth. Phsiological stages in tooth development are: 1.initiation 2.proliferation 3.histodifferentiation 4.morphodifferentiation 5.apposition Initiation
Initiation requires ectomesenchymal-epithelial
interaction It has been demonstrated that dental papilla mesenchyme can induce tooth epithelium & even non-tooth epithelium to form enamel. Lack of initiation:Partial or complete anodontia. Abnormal initiation:single or multiple supernumerary teeth. Proliferation
Enhanced proliferative activity ensues at the points
of initiation & results successively in the bud,cap & bell stages of the odontogenic organ. Proliferative growth causes regular changes in the size & proportions of the growing tooth germ. Histodifferentiation
Succeeds the proliferative stage.
The formative cells of the tooth germs developing during the proliferative stage undergo definite morphologic as well as functional changes & acquire their functional asignment(the appositional growth potential) The cells differentiate & give up their capacity to multiply. This phase reaches its highest development in the bell stage of the enamel organ ,just preceding the formation & apposition of dentin & enamel. The differentiation of epithelial cells preceeds & is essential to the differentiation of the odontoblasts & the initiation of dentin formation. In vitamin A deficiency the ameloblast fail to differentiate properly. Consequently ,their organizing influence on the adjacent mesenchymal cells is disturbed ,& atypical dentin ,known as osteodentin ,is formed. Morphodifferentiation
The relative size of future tooth is establish by
morphodifferentiation,that is by differential growth. Morphodifferentiation is therefore impossible without proliferation The advanced bell stage marks not only active histodifferentiation but also an important stage of morphodifferentiation in the crown ,outlining the future DEJ. The DEJ & DCJ ,which are different & characteristic for each type of tooth acts as blueprint pattern. In confirmity with this pattern the ameloblast,odontoblast & cementoblast deposit enamel,dentin & cementum,& thus give the completed tooth it’s characteristic form & size. Disturbance: affect the form & size of tooth without impairing the function of ameloblast or odontoblast. Results in supernumerary cusp or root ,twinning,loss of cusp or root. “Hutchinson’s incisor” Apposition
Apposition is deposition of hard dental structures
occur in a layer like manner. Appositional growth is characterized by regular & rhythmic deposition of extracellular matrix,which is of itself incapable of further growth. Defects results in “Enamel hypoplasia” “Hypocalcification”. Tooth Type determination
The determination of specific tooth types at their
correct positions in the jaws is referred to as patterning of the dentition. Although in some animals are all the same shape(homodent),in most mammals they are different (heterodont),falling into three families:incisiform,caniniform,&molariform. Two hypothetical models have been proposed to explain how these different shapes are determined 1.Field model theory 2. Clone model theory Field model: Proposes that the factors responsible for tooth shape reside with in the ectomesenchyme in distinct but graded fields for each tooth family. The fact that each of the fields expresses differing combinations of patterning homeobox genes supports this theory. The homeobox genes Msx-1, Msx-2, Dlx-2, and Barx-1 expressed in mandibular mouse ectomesenchyme associated with differing tooth families. Clone model:Proposes that each tooth class is derived from a clone of ectomesenchymal cells programmed by epithelium to produce teeth of a given pattern. Clone theory. A, The molar clone has induced the dental lamina to begin tooth development. At its posterior border the clone and dental lamina grow posteriorly by means of the progress zone. B, When a clone reaches the critical size, a tooth bud is initiated at its center. A zone of inhibition surrounds the tooth bud, and the next tooth bud, C, is not initiated until the progress zone of the clone has escaped its influence