Structure and Properties of Enamel and Dentin

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Structure and properties of enamel


and dentin
V. P. T H O M P S O N, NYU College of Dentistry, USA and
N. R . F. A . S I LVA , Federal University of Minas Gerais, Brazil
DOI: 10.1533/9780857096432.1.3

Abstract: This chapter addresses the mineralized tissues of teeth –


enamel and dentin – and how they develop into structural components
with unique physical properties. Tooth structure includes an epithelium-
derived outer shell of enamel that is highly mineralized, hard, stiff and
wear resistant. This is supported both mechanically and biochemically by
a mesenchyme-derived dentin, which is vital, less mineralized, softer and
more compliant. The dentin is maintained by the dental pulp, which is
cellular and innervated, and has a vascular plexus.

Key words: dentin, enamel, mechanical properties of tooth structure,


mineralized tissues.

1.1 Introduction
Much is known about teeth and their structure. Teeth have long been
studied by paleontologists, since they degrade much more slowly than bone;
in fact, they are the source of our primary knowledge of many ancient
species. Nonetheless our understanding of their intriguing structure is still
incomplete. Human teeth are generally representative, with an epithelium-
derived outer shell of enamel that is highly mineralized, hard, stiff and wear
resistant. The enamel is supported both mechanically and biochemically by
a mesenchyme-derived dentin, which is vital, less mineralized, softer and
more compliant. Dentin is maintained by the dental pulp, which is cellular
and innervated, and has a vascular plexus. In this chapter we give details
of each of the mineralized tissues and how they develop into structural
components with unique physical properties.

1.2 Enamel
1.2.1 Development
Tooth enamel is the hardest tissue in the body, with a hardness comparable
to that of window glass, and is highly fatigue- and wear-resistant. Human
enamel is laid down by cells in a programmed temporal and spatial sequence

3
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4 Non-metallic biomaterials for tooth repair and replacement

to provide the overall shape of the tooth. The cells that make enamel
develop from the invagination of epithelial tissue during fetal development.
In what is known, because of its shape, as the ‘bell stage’ of tooth develop-
ment (ca. 14th week of intrauterine life), the epithelial cells on the inside
of the bell align with a concentration of mesenchyme cells in what appears
to be a one-to-one relationship. More accurately the latter are ‘ectomesen-
chyme’ cells, as the first branchial arch, whose ectodermal cells migrates
into the mesenchyme in the area of the developing jaws (Nanci, 2008).
During this alignment an extracellular collagen network is created that
extends from the epithelial cells to the mesenchyme cells. The epithelial
cells begin to elongate and transform into ameloblasts, and the mesenchyme
cells transform into odontoblasts (Nanci, 2008). The elongation of the
ameloblasts when compared with the odontoblasts leads to pulling on the
collagen network formed between the two, creating a local puckering of
this structure that will become the dentin–enamel junction (DEJ). Seen in
cross-section the DEJ appears as scalloped, but viewed in three dimensions
(3-D), when the enamel has been dissolved, the circular ridges and pits of
the DEJ structure become apparent. The gene expression controlling this
process is not fully understood, but a large number of genes involved in
tooth development have been identified (Nieminen, 2007).

1.2.2 Enamel prisms


The ameloblasts are arranged in a close, overlapping array. Each cell has a
tail that extends between its neighbors (see Fig. 1.1), so that if observed
from above the DEJ, they interdigitate.
Once aligned with their neighbors, the ameloblasts begin to mature and
to lay down the enamel structure. The maturation of ameloblasts starts from
what will become the cusp tip or the incisal edge of the tooth (but at this
stage is the inner top of the bell) and proceeds apically. The last enamel to
begin formation will be that closest to the cement–enamel junction (CEJ).
The ameloblast at its terminal end (nearest to the DEJ) takes on a ‘brush
border’ appearance and begins to excrete proteins, in particular amelo-
genins; these are the template molecules for the nucleation of calcium
phosphate to form, with maturation, ribbons of dense hydroxyapatite (HA).
In this process each ameloblast will create one enamel prism of approxi-
mately 5 μm in diameter, which is also referred to as an ‘enamel rod’ (Fig.
1.2). Individual prisms are currently thought to extend from the DEJ to the
enamel surface through various paths and not to change diameter.
Prisms are joined to their neighbors by a thin organic layer referred to
as a ‘prism sheath’. When loaded to the point of cracking, the resultant
cracks preferentially propagate through the protein sheath, going around
and along the prism (Fig. 1.3).

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Structure and properties of enamel and dentin 5

Enamel crystallites

1.1 Ameloblasts arranged next to one another (upper right). Each


cell has a head (dotted black oval) and a tail (dotted black box) that
extends between its neighbors. Observe the discontinuity of the
enamel crystallites. Asterisk shows secondary territories. Each arrow
in the upper right denotes a sectioning plane through the enamel.
Each arrow points to the diagram depicting the microscopic view of
that sectioning plane in the enamel. Image modified from Boyde
(1989).

5 μm

1.2 Scanning electron micrograph of enamel rods: alignment of


enamel prisms observed when the enamel surface is etched by acid.

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6 Non-metallic biomaterials for tooth repair and replacement

100 μm

1.3 Cracks (crenellations) propagating through the protein sheath


going around and along the prisms following Vickers indentation.

The tensile strength of enamel is lower when loaded perpendicular to the


prism direction (11.4 ± 6.3 MPa) than when it is when loaded parallel (24.7
± 9.6 MPa) (Carvalho et al., 2000). When acid etched, the shear bond of
adhesive applied end-on to the prism direction (enamel surface) is approxi-
mately 40% higher than when the adhesive is applied parallel to the enamel
prism direction (Ikeda et al., 2002). However, self-etch adhesives, which
do not employ a separate etching step, do not result in a significant differ-
ence in bond strength relative to enamel prism orientation (Shimada and
Tagami, 2003).
The laying down of enamel by the ameloblasts proceeds at a rate of about
4 μm per day (Dean, 1998). If an ameloblast were to migrate directly to the
enamel surface, the fastest it could reach the outer dimension of a 1.2-mm-
thick enamel cusp would be (1200/4 =) 300 days, but we note that amelo-
blasts do not proceed directly radially from the DEJ to the surface (as
discussed below), so much more time is necessary to develop enamel for
permanent teeth. Molar enamel thickness varies by cusp from 1.2–1.7 mm
(Mahoney, 2008), increases from the first molar to the third (Grine, 2005)
and is generally slightly thicker for females (Smith et al., 2006). The enamel
thickness on the facial or incisal of a central incisor is approximately
1.3 mm (Shillingburg Jr. and Grace, 1973). Once ameloblasts reach the outer
extent of the enamel they transform to a more cuboidal shape and die. What
signaling controls this process is not known. The calcification of the devel-
oping enamel prism occurs gradually and continues for a some time even
after the tooth erupts into the mouth. This makes newly erupted teeth sensi-
tive to decalcification and caries for more than a year.

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Structure and properties of enamel and dentin 7

1.4 Variation in the width of the daily growth increment and in the
density of calcification. Note striae of Retzius (double headed arrows)
and the instantaneous forming front (white solid arrows) and enamel
prism orientation and direction of growth (white dotted arrow).

Enamel growth periods are seen via structural features in the enamel.
Ameloblasts mature in layers or fronts from the cusp toward the DEJ,
resulting in layers called ‘striae of Retzius’. These appear at the external
surface of the tooth as ‘perikymata’, more pronounced layers in the cervical
enamel surface. Between striae in humans, there are 8–9-day growth incre-
ments designated as the ‘repeat interval’ (Bromage et al., 2011). Within the
repeat interval of enamel there are ‘rhythms’, seen as variation in the width
of the daily growth increment and in the density of calcification (Fig. 1.4).
These same rhythms are seen in the lamellae of bone (Bromage et al., 2011).
The properties of enamel prisms and how these properties change with
prism orientation have been studied extensively since the advent of nano-
indentation techniques (Carvalho et al., 2000; He and Swain, 2007, 2009;
Guidoni et al., 2008). Enamel is hardest along the central axis of the enamel
prism because of the alignment of the HA crystallites in this direction. The
crystallite direction changes across the prism diameter (face) as well as in
the transition between the tail and the body of the rod (Jeng et al., 2011)
(Fig. 1.5). Nanoindentation across the enamel allows the mapping of hard-
ness and elastic modulus, with enamel shown to be both harder and of
higher modulus at the outer surface of the tooth (Angker et al., 2004; Xie
et al., 2009). Higher hardness and modulus are the basis for the wear resis-
tance of the enamel surface. The higher hardness is likely to be related to
the parallel alignment of the enamel prisms over large areas of the outer
enamel surface, an alignment observed when the enamel surface is etched
by acid (see Fig. 1.2). Enamel changes with age, becoming harder at the
surface but not at the DEJ (Park et al., 2008).

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8 Non-metallic biomaterials for tooth repair and replacement

Crystallites

1.5 Crystallite direction changes across the prism diameter (face) as


well as in the transition between the tail and the body of the rod
(black arrows). Image modified from Avery and Chiego (2005).

The unique abilities of enamel, based upon its structure, to withstand


cracking and resist fatigue are in part attributed to the complex pattern
made by the ameloblasts as they traverse and fill the space between the
dentin and enamel surface. A finite number of ameloblasts must each con-
tribute to this process and groups of them seem to act in unison to create
what is known as ‘enamel prism decussation’ (crossing of groups of rods).
Decussation leads to the Hunter–Schreger bands (HSB) seen in sections of
enamel viewed with the light microscope (Lynch et al., 2011). In humans
such decussation is derived from bundles of what are thought to be 50–100
prisms that follow a complex path from the DEJ to the surface in what is
known as ‘multiserial patterning’. These prism groups may be those associ-
ated with a scallop on the DEJ. They can be seen fanning out from the DEJ
in incremental, stacked planes proceeding apically from the cusp tip or the
incisal edge, each plane oriented approximately parallel to the occlusal
plane of the tooth. In each plane the ameloblast groups grow outward at
roughly a 40-degree angle to the radial direction, with one plane orienting
left and the maturing plane below it orienting right. From incisal to gingival
in each plane there are several prism groups. This thickness of the bands
can be seen in a buccal to lingual vertical section of a molar cusp taken in
polarized light (Fig. 1.6). Note that the decussation plane also changes direc-
tion occlusally and gingivally as it proceeds outward. Near the cusp tip or

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Structure and properties of enamel and dentin 9

1.6 Enamel decussation plane changes direction occlusally and


gingivally as it proceeds outwards.

incisal edge the decussation pattern becomes more complex as the amelo-
blast fronts proceed to fill the space. When sectioned this complex pattern
beneath the cusp tip is referred to as ‘gnarled enamel’ (Dean, 1998).
The incisal or occlusal plane of decussation provides enamel with the
ability to resist cracking in this overall direction, as a crack must run a very
long distance to traverse the structure (Bajaj and Arola, 2009a; Myoung
et al., 2009; Bechtle et al., 2010; Ivancik et al., 2011). This is not the case in
the incisal or occlusal to gingival direction. Teeth often show vertical cracks
in enamel without consequence but rarely horizontal cracks, as the latter
are quite detrimental (Lee et al., 2011). Researchers have been investigating
the ‘fracture toughness’ of enamel, that is, the energy necessary to propa-
gate a crack (Bajaj et al., 2008; Bajaj and Arola, 2009a; Ivancik et al., 2011).
Using very small sections of enamel to make compact tension and fracture
toughness coupons, they have shown that the fracture toughness from
surface inward or from the DEJ outward increases by an order of magni-
tude as the crack extends from either surface. They attribute this impressive
behavior to enamel decussation (Bajaj and Arola, 2009b). Testing of whole
teeth shows high resistance to cracking, with exceptional resistance in the
occlusal as opposed to the vertical plane (Chai et al. 2009, 2011).

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10 Non-metallic biomaterials for tooth repair and replacement

Although enamel is hard and wear resistant with its high HA content
(> 95% by weight), it is still a hydrated tissue and chemicals can diffuse
through the structure surprisingly rapidly. Once through the enamel the
chemical species makes rapid access to the pulp via the dentinal tubules
containing the odontoblastic process, making dentin highly permeable. In a
cat canine model Lucifer Yellow dye can penetrate from the enamel surface
to the odontoblasts in the pulp within 30 min (Ikeda and Suda, 2006). In
extracted teeth subjected to bleaching agents, peroxide is present in the
pulp within 30 min of external application (Gokay et al., 2000). Given the
ready permeability of enamel we can hypothesize that microscopic cracks
on the surface and at the DEJ may be able to heal through remineralization
from saliva or from dentin interstitial fluid, respectively.

1.3 Dentin–enamel junction (DEJ)


As noted earlier, the DEJ is the interphase between enamel and dentin,
initially formed by the alignment of ameloblast and odontoblast during the
bell stage of tooth development. It is approximately 60–100 μm in width
and is a ‘graded structure’, in that the elastic modulus makes a nearly linear
transition from the enamel (∼70 GPa for inner enamel) to that of dentin
(∼15 GPa) (Marshall et al., 2001; Chan et al., 2011). The change in the
calcium and phosphate content from the enamel to the dentin is thought
to be responsible for this gradient in modulus. A graded structure serves to
lower tensile stresses substantially at the interface of a brittle material with
one of lower elastic modulus, resulting in increased strength and fatigue
resistance (Zhang and Ma, 2009; Ren et al., 2011). The DEJ gradient moves
the highest tensile stresses into the bulk of the enamel during function and
reduces those at the interface by nearly 50% (Huang et al., 2007).
A graded interphase has also been identified between cementum and
dentin (Ho et al., 2004). Dentin mineral content changes from the DEJ
toward the pulp, as noted by change in HA particle size (Marten et al.,
2010).

1.4 Dentin
The ectomesenchyme cells that become odontoblasts align with the
ectodermal cap cells that become ameloblasts (Nanci, 2008). While the
ameloblasts are elongating, the odontoblasts are already beginning to
produce the collagen network that becomes the DEJ; they then make the
transition to elaboration of the more complex collagen and proteoglycan
structure of dentin. The elaboration of the dentin structure and its following
calcification proceed inwards at an initial rate of ∼2.8 μm per day. This rate
slows as the odontoblasts approach the pulp space and slows further when

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Structure and properties of enamel and dentin 11

2 μm AF

1.7 Lateral extension of dentinal tubule (white dotted arrow) of an


etched dentin substrate.

root formation and tooth eruption are complete. Dentin continues to grow
inwards for the life of the individual and leaves a record of growth-altering
events. This is termed ‘secondary dentin formation’. Dentin is also dynamic
in that it can respond to insults to the enamel, such as caries or excessive
wear, and lay down additional dentin, referred to as ‘reactionary dentin’ or
‘tertiary dentin’ (Bjorndal and Darvann, 1999; Bjorndal, 2001). Wear of root
surfaces can also lead to laying down of reactionary dentin (Nanci, 2008).
Dentin has a structure with tubules that course from the DEJ to the pulp
radially inwards, with a broad S shape when the tooth is sectioned axially.
Within the tubules are cellular processes extending from the odontoblasts
that line the pulp. The tubules have smaller lateral extensions along their
length that communicate with neighboring tubules, creating a communica-
tion and interstitial fluid network that maintains the dentin. These lateral
extensions can be seen in sections of etched dentin (Fig. 1.7).
The collagen structure of dentin is complex, with the collagen oriented
in helical-like structures forming tubules but then changing to a more radial
orientation in the plane perpendicular to the tubule direction. There are
proteogylcans aligned along collagen fibers and these play a role in miner-
alization and physical properties (Chiu et al., 2012). Calcification of dentin
starts with nucleation in the gap space between collagen strands and pro-
ceeds outwards expanding in the direction of the fibers, forming elongated
crystals of HA that are anisotropically oriented to withstand loading

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12 Non-metallic biomaterials for tooth repair and replacement

(a) (b)

1.8 (a) Near DEJ dentin tubules that are widely spaced and about
0.8–1.2 μm in diameter, with a distance between tubules of nearly
10 μm. (b) Tubule density increases to approximately 40 000 per mm2
near the pulp.

(Marten et al., 2010). The dentin around the tubules is more highly mineral-
ized; this zone of mineralization, approximately the thickness of the tubule
diameter, is called the ‘peritubular dentin’. Outside this zone the mineral
content is lower; these regions comprise the ‘intertubular dentin’.
Near the DEJ, the dentin that forms has tubules that are widely spaced
and about 0.8–1.2 μm in diameter, with a distance between tubules of nearly
10 μm (Fig. 1.8(a)). This gives a tubule density of 17 000–20 000 per mm2.
With the radial orientation of the tubules, tubule density increases to
∼40 000 per mm2 near the pulp (Pashley et al., 1985) (Fig. 1.8(b)).
Inner dentin has a reduced amount of intertubular dentin, but this does
not lead to an increase in hardness: the overall mineral content changes
from the DEJ toward pulp as the HA particle size decreases with depth
(Pashley et al., 1985; Marten et al., 2010). Note that dentin that has been
etched and then dried has a collapsed collagen layer that appears as a gel
(Fig. 1.9). Contrast this with the visible collagen seen in freeze-dried etched
dentin (Fig. 1.10).
The properties of dentin have been studied to determine its strength with
orientation as well as its fracture toughness. Using microtensile specimens
Gianni and others (Giannini et al., 2004) have shown that the tensile strength
of dentin perpendicular to the tubule direction is 62 GPa near the DEJ and
reduces to ∼34 GPa near the pulp. This group also related the ability to
bond adhesively to dentin to the area of the intertubular dentin (Giannini
et al., 2001) and, inversely, to the tubule density. The hardness of dentin on
a macroscopic scale (Knoop or Vickers indentation) is isotropic perpen-
dicular or parallel to the tubule direction at the same relative depth in the
enamel (Pashley et al., 1985). Hardness is reduced with depth in dentin
(Hosoya and Marshall, 2004) and varies from buccal to lingual (Brauer
et al., 2011). Radicular intertubular dentin has a reduced elastic modulus
and hardness compared to coronal intertubular dentin (Inoue et al., 2009).

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Structure and properties of enamel and dentin 13

1.9 Scanning electron image of a dentin substrate that has been


etched and then dried, showing a collapsed collagen layer that
appears as a gel.

5 μm

1.10 Scanning electron image of collagen seen in a freeze-dried


etched dentin surface (courtesy of Dr Jorge Perdigão).

Dentin can also be considered to be a graded structure, given that it changes


properties with location (Tesch et al., 2001).
Dentin toughness has been studied to understand the mechanisms that
limit crack extension (Imbeni et al., 2003; Kruzic et al., 2003; Nalla et al.,
2003). In their review, Nalla and others (Nalla, et al., 2003) show that crack
bridging and the formation of daughter cracks are significant mechanisms
for the dissipation of crack energy, so that the process leading to the tough-
ness of dentin is similar to that of fracture toughness in bone. Replacing the

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14 Non-metallic biomaterials for tooth repair and replacement

Table 1.1 Physical properties of enamel and dentin compared to other


dental materials

Tooth Modulus (GPa) Hardness (GPa) Toughness (MPa m1/2)

Enamel 94 3.2 0.8


Dentin 16 0.6 3.1
Resin cement 4–5 0.3 1.1
Composites (Z 100) 18 – 1.3
Glass-ceramic 67–96 3.4–6.3 100–420

water in dentin with less polar solvents, such as ethanol, increases the
fracture toughness of dentin (Nalla et al., 2005, 2006). However, dentin
fracture toughness is reduced with age (Kinney et al. 2005; Nazari et al.
2009), which may help to explain the significant increase in cusp fracture
of posterior teeth with age, in particular those that have been restored.
Restoration often leads to volumes of dentin where the dentin tubules
have been cut and thus can no longer supply interstitial fluid minerals to
the dentin and associated DEJ and enamel. Table 1.1 presents physical
properties of enamel and dentin compared to commonly use restorative
materials.
Dentin is dynamic in that it reacts to the caries process with a low perme-
ability zone and can remineralize caries-affected areas if the caries is sealed
from the oral environment (ten Cate, 2001, 2008). Caries established in
dentin is characterized as comprising a bacterial infected layer adjacent to
the enamel and, beneath this, an acid-altered demineralized zone desig-
nated as ‘affected dentin’. This demineralized zone, detected by dyes, is
inaccurately perceived by clinicians to be ‘infected dentin’ (Boston and
Liao, 2004). Within the affected dentin near the bacterial front the acid
attack has dissolved most of the mineral, but beneath this is a zone where
the acid attack is actively dissolving the HA. This dissolution yields a sig-
nificant concentration of calcium and phosphate, leading to precipitation of
an acid-resistant compound, whitlockite, in the dentinal tubules (Daculsi
et al., 1987). Histologically this is seen in thin section and identified as
the ‘transparent zone’. This tubule precipitate lowers the dentin permeabil-
ity (Pashley et al., 1991), allowing excavation of infected and overlaying
affected dentin without anesthetic being required (Boston, 2003; Allen
et al., 2005).
Exposed dentin root surfaces of teeth may become worn through abra-
sion, erosion, or a combination of both, perhaps accelerated by occlusal
stress (Gallien et al., 1994; Terry et al., 2003; Pecie et al., 2011). Often such
dentin has a smooth, hypermineralized surface called ‘sclerotic dentin’ (Aw
et al., 2002). The low-permeability zone in affected dentin is sometimes

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Structure and properties of enamel and dentin 15

referred to as ‘sclerotic dentin’, but this is a misnomer: the calcification is


in the tubules, whereas the surrounding dentin may be undergoing decalci-
fication. Root surface sclerotic dentin in non-carious cervical lesions pres-
ents a challenge for dentin bonding agent procedures (Yoshiyama et al.,
1996; Marshall et al., 2000), as most are evaluated on normal or caries-
affected dentin.
Now well established but not appreciated clinically is the ability of caries
to be arrested and undergo varying degrees of remineralization if sealed
from the oral environment (Carvalho et al., 1998; Thompson and Kaim,
2005; ten Cate, 2008; Alves et al., 2010; Bjorndal, 2011). Quantification
of this process has recently been observed with and without use of a
calcification-promoting liner containing amorphous calcium phosphate
compounds (Bresciani et al., 2010; Peters et al., 2010).

1.5 Conclusion
Teeth are unique biological structures that can last a lifetime in service.
Nature, using a cellular approach, has constructed a fatigue- and damage-
resistant structure, and to some extent a self-healing one, that is now only
being approached in performance by ceramic and resin-based composite
formulations. In studying the structure of enamel and dentin we may be
provided with clues about the design and development of new materials
with broad-ranging applications.

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Structure and properties of enamel and dentin 17

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