Life Cycle of ameloblasts

Dr Talaya Zahid
AMELOGENESIS
Stages
• Presecretory
Morphogenesis
Histodifferentiation

• Secretory
Initial secretory without Tomes processes
Late Secretory with Tomes processes

• Maturation
Ruffle-ended Ameloblast
Smooth-ended ameloblast
Presecretory Stage

Divided into
• Morphogenic phase
• Differentiation phase
Morphogenic phase

• During the bell stage of tooth development, the shape of the crown is
determined.
• At this stage, dentin is not mineralized.
• The cells of the inner enamel epithelium still can undergo mitotic
division, throughout the bell initially and eventually limited to the
cervical portion of the tooth.
• These cells are cuboidal or low columnar, with large, centrally
located nuclei
• Poorly developed Golgi elements in the proximal portion of the cells
Differentiation phase

• Inner enamel epithelium cells differentiate into ameloblasts

• they elongate and their nuclei shift proximally toward the stratum
intermedium.
• During mentle dentin formation basal lamina supporting these cells is
disintegrated
• Golgi apparatus and ER increases in number and golgi shifts distally
contd..

• At distal extremity of the cell, the ameloblast differentiated into a

body and a distal extension called Tomes’ process, against which
enamel forms(POLARIZED CELL)
• Ameloblast start secreting before basal lamina is disrupted which is
seperating preameloblasts and preodontoblasts
• Preameloblast also secret dentine sialoprotein
• At later stages reciprocal expression of opposing matrix proteins and
production of typical mesenchymal proteins by enamel organ derived
cells are evident
• ameloblasts are aligned closely with each other with junctional
complexes
• Fine actin-containing filaments radiate from the junctional
complexes into the cytoplasm of the ameloblasts distinguishing as
proximal and distal terminal webs
Secretory stage

• ameloblasts reflects their intense synthetic and secretory activity

• Golgi bodies surrounded by cisternae of ER is present
• mRNA is translated by ribosomes on RER
• Proteins are synthesized and translocated into cisternae
• Proteins are packed by golgi bodies as membrane bounded secretory
granules
• These proteins migrate into tomes process(continuous secretion)
• Tomes process comprises only a proximal portion when enamel
formation is just at begining
• secretory granules product is released against the newly formed
mantle dentin to form enamel(having no rods)
• This is how first hydroxyapatite crystal formed
• Initial enamel layer is formed
• Ameloblasts migrate away from the dentin surface and develop the
distal portion of Tomes process as an outgrowth of the proximal
portion
• distal portion of process penetrates into and interdigitates with the
enamel beyond the initial layer
• Rod and interrods are formed by ameloblast and its processes
• Enamel proteins are released extracellularly can be identified by the
presence of abundant membrane infolding
• Secretion from the first site on the proximal part of the process
results in the formation of enamel partitions that delimit a pit
• In this pit there resides the distal portion of Tomes process.
• Walls of pit formed by interrod
• Secretion from the second site distal portion of tomes process provide
matrix for enamel rod
• Interrod formation is completed before enamel rod formation
• Rod and interrod enamel differ only in the orientation of their
crystallites
• Distal portion of tomes process lengthens as enamel thickens
• Gets thinner as the rod growing in diameter presses it against the wall
of the interrod cavity
• Distal tomes process sqeezed out
• This creats a narrow space along most of the circumference between
rod and interrod enamel that fills with organic material to form rod
sheath
• Rod crystals formed in relation to the secretory surface directly
against the interrod partition
• shape of the distal portion of Tome’s process is altered when enamel
outer layer is formed and its orientation is also changed
• Thus, enamel rods in the outer third of the enamel layer have a
slightly different profile and have a more rectilinear trajectory
• Ameloblasts are now shorter by losing its distal portion
• So without distal portion enamel having same appearance without
rods as during initial lay down of enamel(as rods form in relation to
the distal portion of Tome’s process)
• Making prismatic sandwhich btw aprismatic enamel
Matrix proteins

• Amelogenin (in early maturation stage)

• Ameloblastin (expressed untill later stages)
Division of maturation phase

• Early maturation phase

• Transitional phase
• Maturation proper
MATURATION STAGE(early)

• Before the tooth erupts in the oral cavity, enamel hardens.

• This change in physicochemical properties results from growth in
width and thickness of preexisting crystals seeded during the
formative phase of amelogenesis and not because additional crystals
are created de novo
• Crystal growth during the maturation stage occurs at the
expense of matrix proteins and enamel fluid that are
largely absent from mature enamel
• Up to about two thirds of the formation time can be
occupied by the maturation stage
• Maturation stage ameloblasts seem to carry out small,
repeated developmental increments with a cumulative
effect of great change.
• Although maturation stage ameloblasts generally are
referred to as postsecretory cells, they still synthesize
and secrete proteins
• These ameloblasts still exhibit a prominent Golgi complex, a
structural feature consistent with such activity

• Although amelogenin signals are found only in the early maturation

stage, those for ameloblastin continue to be expressed until much
later
Transitional phase

• Full thickness of immature enamel has formed

• Ameloblasts show morphological changes
Events in transitional phase

• Reduction of ameloblast height

• Decrease in organelle content
• Apoptosis of ameloblasts
(25% cell die during transitional phase and 25% die as enamel matures)
• Apoptosis can also be seen in enamel knot
(cells die to permit orderly morphogenesis at specific times)
Cell death

• Cells die by two ways

1. Necrosis
2. apoptosis
Regulator proteins for cell death

• Bcl-2 family proteins antiapoptotic and proapoptotic proteins

• Prateinases (caspases)
 MATURATION STAGE(proper)
• Next the principal activity of ameloblasts is the bulk
removal of water and organic material from the
enamel to allow introduction of additional inorganic
material

• The most visually dramatic activity of these cells is

modulation i.e. the cyclic creation, loss, and recreation
of a highly invaginated ruffle-ended apical surface
(the cells alternate between possessing a ruffled
border [ruffle-ended] or a smooth border [smooth-
ended]
 MATURATION STAGE(proper)
• Modulation can be visualized by special stains and
occurs in waves traveling across the crown of a
developing tooth from least mature regions to most
mature regions of the enamel (e.g., in an apical-
incisal direction in continuously erupting teeth and
cervical-incisal [occlusal] direction in teeth of limited
eruption).
• They appear to be related to maintaining an
environment that allows accretion of mineral content
and loss of organic matrix, in part through alterations
in permeability of the enamel organ
FIGURE 7-14 Schematic
representation of the various
functional stages in the life cycle of
ameloblasts as would occur in a
human tooth.
1, Morphogenetic stage; 2,
histodifferentiation stage; 3, initial
secretory stage (no Tomes’
process); 4, secretory stage (Tomes’
process); 5,
ruffle-ended ameloblast of the
maturative stage; 6, smooth-ended
ameloblast of the maturative stage;
7, protective stage.
MATURATION STAGE(proper)

• Ruffle-ended ameloblasts show considerable

endocytotic activity and contain numerous
lysosomes, calcium-binding proteins, and
membrane-associated calcium-
adenosinetriphosphatases that appear to promote the
pumping of calcium ions into the maturing enamel.
• Smooth-ended ameloblasts, however, leak small
proteins and other molecules, show little
endocytotic activity, and have almost no membrane
calcium-adenosinetriphosphatase activity.
MATURATION STAGE(proper)

• The calcium ions required for active crystal growth

pass through the ruffle ended ameloblasts (because
their distal junctions are tight) but along the sides of
the more leaky smooth-ended ameloblasts.
• Regarding the withdrawal of organic matrix from
maturing enamel is attributed largely to the action of
bulk-degrading enzymes that act extracellularly to
digest the various matrix proteins into fragments
small enough to be able to leave the enamel layer
MATURATION STAGE(proper)
• Polypeptide fragments leaving the enamel likely
pass between the leaky distal junctions of smooth-
ended cells and diffuse laterally among the
ameloblasts to be taken up along their basolateral
surfaces.

• When cells become ruffle-ended, because the

proximal junctional complex now in turn becomes
leaky, some of the laterally diffusing peptides could
disperse throughout the papillary layer and perhaps
beyond
MATURATION STAGE(proper)
• Just as ameloblasts complete the transitional phase
and begin the first series of modulation cycles, they
deposit an atypical basal lamina at their now-
flattened apex (no part of Tomes’ process is
recognizable at this stage).
• This interfacial layer adheres to the enamel surface,
and the ameloblasts attach to it by means of
hemidesmosomes
• Basal lamina constitutes contain laminin-332
heterotrimer molecule that is essential for the
formation of hemidesmosomal attachments
 MATURATION STAGE(proper)
• This atypical basal lamina is
known to be rich in
glycoconjugates and to contain
some unique proteins
• Detection of sugar residues
using lectins, here Ricinus
Communis Agglutinin (RCA)
specific for galactose,
indicates that the basal lamina
is rich in glycoconjugates
MATURATION STAGE(proper)

• It likely represents a unique structure both in

composition and function; in addition to an adhesive
role, the presence of highly glycosylated molecules
may confer to this lamina charge-selective property
that could help regulate the movement of material
into and out of the enamel layer

• It is situated such that it could relay to the

ameloblasts information about the status of the
dynamic enamel compartment.
MATURATION STAGE(proper)
• Initially, the cells are involved in establishing the
crown pattern of the tooth (morphogenesis); at this
time they are small and low columnar with
centrally placed nuclei, and they undergo frequent
mitoses.
• Then, the cells undergo morphologic changes, and
they become ameloblasts (histodifferentiation).
• These changes are preparatory to their entering the
next phase, active secretion of the enamel matrix,
wherein they develop a cell extension called the
Tomes’ process.
MATURATION STAGE(proper)

• The secretory stage is followed by a short

transitional phase of cell restructuring leading to
enamel maturation proper, wherein the ameloblasts
exhibit cyclical variations with ruffle- and smooth-
ended borders against the enamel surface; the ruffle-
ended cells allowing incorporation of inorganic
material, the smooth-ended cells permitting exit of
protein fragments and water.
• The final phase is protection of the newly formed
enamel surface until the time of tooth eruption
Summary
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