Digestive System: Oral Cavity

Adam and Eve had many advantages, but the principal one was that
they escaped teething.

—Mark Twain (1835-1910)

Pudd'nhead Wilson, Chapter IV

Objectives for This Exercise

SLIDES FOR THIS LAB: 1203, 1210, 1205, VM58, VM62, 1-8, 85

The oral cavity is the first portion of the digestive

tract, and consists of the mouth and the
structures which are enclosed within it. The
mouth isn't merely a hole in the face. It's a
complex structure with nutritional, respiratory,
and communicative functions in all mammals and
most other groups of animals.

Additionally, it serves as the principal means of manipulating (if you'll

pardon the expression) the environment for those animals which
don't have hands, such as dogs. A dog carries things in his mouth
because he has no other way to do it. (If you have time someday, try
to get a dog to bark when he has something in his mouth. This takes
some work, but the noise he makes when trying to eat and
communicate simultaneously is worth hearing.)

One can get a clue to the nature of an animal's diet by the type and
shape of the teeth, which differ greatly in carnivores and herbivores.

Accessory structures which contribute to the mouth's function

include the tongue, the teeth, and the salivary glands. The tongue and
the teeth will be dealt with below: click here for a discussion of the
structure of salivary glands.
 Oral Epithelium

The oral cavity in

domestic animals is
lined with stratified
squamous epithelium.
Depending on the
nature of the animal's
diet, this may or may
not be keratinized.
Those animals which
eat a great deal of
roughage, such as
ruminants and horses,
usually have a very
heavily keratinized oral
cavity, while those
which eat softer food
show less of it. There is
a considerable amount
of variation in this rule,
both across species
lines and among
individuals in a species.

This is a section
through the wall of the
oral cavity, i.e., a cheek
or lip. It's cut vertically,
and you are looking into
it from the side.

The oral side has a lining of stratified squamous epithelium. Since

this specimen is from a primate, it's not keratinized. The oral cavity
side totally lacks any of the adnexal structures associated with skin:
no hairs, sebaceous or sweat glands, etc. The skin side has these
things, and as external integument, it's always keratinized to some
degree. The sinus hairs are the very large ones associated with tactile
sensation through the vibrissae.

The center of this specimen contains some of the buccal muscles,

strands of skeletal muscle that are part of the muscle sheet
underlying the skin. Here it's acting as the "tunica muscularis" for this
part of the digestive tract. Many rodents, such as the Golden Hamster
(Mesocricetus auratus) have cheek pouches for temporary storage of
food. The cheek pouch muscles are very well developed. The hamster
fills the pouch and then can contract the buccal muscle to dump the
contents whenever he wants.

The place of continuity between the "dry" integument and the "wet"
oral mucosa is located at the top of this section. This sort of
transition point is a mucocutaneous junction. Mucocutaneous
junctions exist in several other sites: the anus, the nostrils, the vulva
and the urethra.

Tongue

Slide 1210
is a section
of a
tongue.
The tongue
in mammals
is an
extremely
muscular
organ
within
whose
substance
there are a
number of
smaller
salivary
glands, and any number of motor and sensory nerve fibers. The bulk
of the tongue is skeletal muscle, arranged in three layers, all at right
angles to each other. This provides for an amazing degree of
flexibility and is vital to vocalization. There may be a considerable
amount of adipose tissue present as well. The orientation of the
muscle layers gives the tongue a distinctive appearance in LM
sections that isn't easily mistaken for anything else.

The image above at the right is a cross section of a mouse tongue.

The snout of the mouse has been cut so that you are looking "into"
the oral cavity, i.e., it's a "frontal" section plane. The top of the image
is the dorsal side, the bottom ventral; the tongue is the large oval
structure in the center.
The space around it is
the oral cavity.

The dorsal surface of the

tongue is covered with
specialized lingual
papillae (see below),
special regions of the
stratified squamous
epithelium that covers
the entire organ. Not
much detail can be see
in this low power image,
but a closer look will
reveal the way the muscle bundles are laid out. In this higher
magnification view, the muscle bundles and their arrangement can be
seen clearly. This organization of muscle bundles is one of the
identifying features of the tongue. One set runs parallel to the screen
from upper left to lower right; a second set runs at a right angle to
these from lower left to upper right. The third group is cut in cross
section, as it runs in and out of the plane of the screen; it's therefore
at right angles to both the other two. The bundles are thus set into X,
Y, and Z axes. The muscle bundles play off against one another, and
by controlling the tension it's possible for the tongue to assume many
shapes.

This requires exquisite neural control, and the tongue has that. It's
common to see nerve fibers running between the bundles, and small
salivary glands here and there (left). The muscles are controlled by
motor fibers from cranial nerves; there are fibers for general touch
sensation (quite a few of them) and those serving the taste buds on
the dorsal surface. Sympathetic and parasympathetic fibers innervate
the salivary glands, and there are ganglia embedded in the substance
of the tongue as part of the "wiring" for the glands. In the image
above right, neurons of one of these ganglia are indicated by the
arrows.

Filiform Papillae
On the tongue's dorsal surface slide 1205 shows numerous filiform
papillae. These are slender conical structures, which project upwards
forming a velvety covering on the tongue. The filiform papilla is one
of 4 types of lingual papillae, and its function is purely mechanical; it
has no sensory structures associated with it, as do the others. In
some species (such as cats and cattle) filiform papillae are extremely
large and rough, and the peculiar rasping effect of a cat's tongue is
due to these papillae. It's also what a cat uses to clean her fur. The
stiff papillae make effective bristles for removing debris. You can also
see some of these on slide 115.

The filiform papillae are by far the most common type, and sometimes
are "swallow tailed" in shape. They're raked backwards. The "spines"
on a cat's tongue are a nice example of this type of papilla, and
they're what gives the tongue its rough feel and velvety appearance.
They have no sensory function. In large animals, such as cows, the
filiform papillae can be very large, up to perhaps 5.0 mm long. When
they get this big, they sometimes have a "core" of CT inside. But in
smaller animals they are pretty much entirely epithelial in nature.

Other Lingual Papillae

The remaining types of lingual papillae are sensory in function. They

include: the fungiform papillae, which are less common than the
filiforms; the vallate (or circumvallate) the largest and most
prominent; and the foliate papillae, which are most easily seen in the
Order Lagomorpha (rabbits, hares, and conies).

Vallate Papillae

A vallate papilla is present on slide 1210. This type of papilla is set

into a deep pocket in the tongue's surface, and anchored at the
bottom by a short broad stalk. It doesn't protrude above the general
level of the surface by much, but it's surrounded by a deep "moat"
into which some of the lingual salivary glands secrete. Vallate
papillae are easily visible with the naked eye in most animals. They're
paired and located near the back of the tongue. They're the least
numerous type. This very large structure has a core of connective
tissue (CT) and nerve fibers are often seen in it, serving the taste
buds.

Taste buds (TB) are present on the sides of the papilla, and also
frequently on the side of the moat. The small lingual salivary glands
(SG) that empty into the moat keep the taste buds ready for re-use.
There are two types of these lingual salivary glands: serous and
mucous in nature. The secretions of these glands are wash the
chemoreceptor sites of the taste buds free of molecules, so that new
sensations can be detected. These aren't the only salivary glands in
the tongue, by the way; there are small ones scattered throughout the
muscle bundles.

Vallate papillae usually show taste buds, seen as well defined, lightly
staining areas which enclose several banana shaped sensory
transducer cells. You should see some on the undersides of the
papilla proper, and you may also see some on the tongue side of the
"moat." Taste buds are discrete and well encapsulated by delicate
CT, which can be demonstrated with special stains.

The sensory cells of the buds are located below the level of the
epithelium, and they communicate with the outside via a so-called
taste pore, a break in the epithelium. Through this pore project the
microvilli or "taste hairs" on the tops of the sensory cells. The taste
bud is yet another example of a peripheral transducer structure
whose function is to convert some physical phenomenon into a
nervous signal. In this case the phenomenon is the binding of tastant
molecules to the membrane of the transducer cells. Binding causes
charge changes on the cells that are detected by neural fibers
wrapped around them.

Not a whole lot is understood about the phenomenon of taste.

Traditionally there are said to be four basic taste sensations: sweet,
salt, sour, and bitter, but recent research casts some doubt on this
simplistic explanation. Specific regions of the tongue do have specific
associations with these sensations: the data is incomplete, but it's
been shown that generally the tip of the tongue has receptors
for sweet and salty sensation, and sour and bitter receptors lie near
the back. There are no morphological differences that correlate with
functional ones, however, and you can't tell by looking at a taste bud
what kind it is.

It's thought that the different taste sensations are located in and
associated with specific cell types in the taste buds, but there are also
indications that some cells may respond to more than one taste
stimulation. Depending on where they're located, the taste buds may
be innervated by fibers from cranial nerves VII (facial), IX
(glossopharyngeal) or X (vagal). This is separate from the "general"
sensation; that is, tactile perception is not connected through the
same nerve circuits. A branch of cranial nerve V (trigeminal) handles
this input.

Taste sensation is found in all vertebrates, and a similar

chemoreception ability in most if not all invertebrates. The more or
less universal distribution of chemoreception makes it clear that this
sense has very high survival value.

The receptor cells, regardless of type, are more or less banana

shaped, and at the apex of each is a small microvillus, called a "taste
hair." (This is a misnomer because they aren't "hairs" in any real
sense, but we're stuck with this archaic name.) The "taste hairs"
collectively come to form a bundle that's exposed to the world via a
small opening in the overlying epithelium, the taste pore. The "hairs"
are the actual site of chemoreception; the plasma membrane that
covers them contains receptor sites for the different sensations.

Integration of the chemoreception signals takes place in the central

nervous system. The transducers respond, but the brain actually
"tastes" by interpreting the responses. To a great extent, perception
of tastes as "good" or "bad" is learned behavior. In some cultures,
grilled meat is considered delectable, but in others,
nauseating. Similarly, I have lived in places where rotten eggs are
regarded as a delicacy, but my cultural perceptions prevented me
from eating them. Many animals (humans included) have to learn what
foods are edible, and one of the clues they use is taste perception.
 Fungiform and Foliate Papillae

The fungiform papillae, as the name implies, are mushroom shaped

structures. They rise above the general level of the filiform papillae,
and they usually have taste buds. The foliate papilla is best seen in
rabbits. The term means "leaflike" as in the leaves of a book, and
these are seen in the demonstration as if you were viewing the top
edge of a page of paper. They are set in the sides of the tongue, and
you are looking at them from the dorsal surface. Both fungiform and
foliate papillae carry taste buds.

The mushroom-shaped fungiform papilla is nicely illustrated here in

diagram and in an actual
specimen. Fungiform
papillae are far less
numerous than the
filiform type, and
scattered in
among them. They're
not nearly as large as
the vallate papillae, but
they are large enough to
have a core of CT. The
fungiform papillae stick
up above the
surrounding filiforms,
and here you can see the relationship pretty well. A swallow-tailed
filiform papilla flanks this fungiform one on either side.

Here's a group of foliate papillae. The word "foliate" means "leaflike"

and in this case the reference is to the "leaves" of a book. The
orientation of this specimen is different from the ones seen before.
You are viewing these papillae from the top; that is, the tongue has
been cut parallel to its long axis, and exposed to view from the dorsal
side. Thus, you're seeing the edges of the foliate papillae as if you'd
seen the top edge of a book.

The papillae themselves are flat-topped and have deep clefts between
them. The clefts are lined with epithelium, and there are taste buds
embedded in them. This type of papilla is most commonly seen in
members of the Order Lagomorpha, i.e, rabbits, hares, pikas, and
conies. Similar papillae are found in the tongue of the Florida
manatee!

Teeth

Sound teeth are one of the more important things in life, and an
abscessed tooth is likely to make the whole world look black to the
individual suffering from it, two-legged or four-legged. Yes, dogs do
get dental caries, and they can suffer from them, though they're less
likely to have them than humans are because of differences in diet
and
tooth
shapes
.

We
feather
less
bipeds
, and
most
of our
pets,
have b
rachyd
ont
teeth,
like the
one
shown
at left.
This is
the
type
found
in
carniv
ores
and
the
"typical" tooth model. (Ruminants and horses have a different type
which won't be considered in this exercise.) You will find a typical
brachydont tooth on slide 85. The tooth on this slide is developing in
a socket of the jawbone. This bony alveolus is part of the skeleton,
but strictly speaking the tooth isn't.

There are three hard materials involved in tooth structure: enamel,

dentin, and cementum. Enamel covers the outside of the tooth, and in
the processing of a tooth for microscopic examination, it's usually
lost. It's a thin, brittle coat that's easily chipped and almost entirely
made of inorganic materials. Dentin comprises the bulk of the tooth's
mass. It's softer than enamel, and underlies it. Cementum is a bone-
like substance with embedded cementocytes, that covers the root of
the tooth, but doesn't
project above the gum
line.

Brachydont teeth are

hollow. There's a
space inside, the pulp
cavity, that contains
living material in the
form of cells, and a
relatively
undifferentiated
primitive form of CT.
The pulp cavity has
blood vessels that run
in and out to serve the
needs of these tissues,
and also nerve fibers.

Since the tooth has to have a blood supply, and since only CT can
make blood vessels, the periodontal ligament serves not only as the
anchoring material, but as the conduit through which blood vessels
and nerves go to and from the pulp cavity. Access to this space is via
the root canal, visible in the image to the right as a channel through
the hard material. When a dentist does a "root canal" on you, what's
happening is the entire pulp cavity is being purged of its contents,
and filled with restorative material. He has to remove the blood
vessels and nerves, which is done by drilling out not only the pulp,
but all the way down through the root canal, and then stuffing
everything full. The presence of the nerve fibers is what makes this
such an exhilarating experience.

Teeth are anchored into the bone of the jaw by a tough,

collagenous periodontal ligament (sometimes called the periodontal
membrane, PM above) that serves as the periosteum for the bony
socket, and attaches firmly to the base of the tooth, where the
cementum (C) is located. Try this: grab one of your teeth and push it,
fairly hard: it will move, just a bit. That's the "give" in the ligament.
Sometimes teeth get loosened by an impact; when this happens after
a while the ligament will repair the damage and the tooth is firmly
seated once more.

Tooth formation is a complicated process, and the tooth's hard

structures are the result of cellular activity, but are in and of
themselves non-cellular, dead material. Primitive connective tissue,
or mesenchyme condenses in the future central region of the tooth, in
what will become the pulp cavity. Inside this region a special
population of cells, the odontoblasts begin the process of making
dentin, by laying down a matrix material and then calcifying it. As
they do so, they move backwards towards the center of the pulp
cavity. The odontoblasts remain active throughout life, so the process
of making dentin continues after eruption. As the animal ages the
pulp cavity slowly diminishes in size.

On the outside surface of the tooth, a similar process takes place. A

population of ameloblasts differentiates, as a row of nicely columnar
cells facing the odontoblasts. Ameloblasts make enamel in a way
that's comparable to the manufacture of dentin. They lay down an
uncalcified matrix first, then harden it, backing away as the enamel
layer is built up. In brachydont teeth, enamel is made only before
eruption. Once the tooth erupts, the ameloblasts die. If enamel is
damaged it can't be renewed.

You see here a typical forming tooth, in situ. An important thing to

remember is that the hard material of the tooth—the enamel and the
underlying dentin—are both formed extracellularly.
The ameloblasts are ectodermally derived, from the oral epithelium in
the embryonic oral cavity. The embryonic origins of the odontoblasts,
the cells which make the dentin, is a matter of some debate, but they
are resident in the site of the future pulp cavity, which is generally
held to be embryonic mesenchyme. Hence, they are probably derived
from mesoderm. There is recent evidence to suggest, however, that
they may be one of the many derivatives of the neural crest cells, and
ectodermal as well.
Odontoblasts and ameloblasts have some similarities, whatever their
origins. Both cell types are tall and columnar in shape. They lay down
precursor materials—pre-dentin and pre-enamel—and as they do so,
"back away" from it. The precursor material is then calcified as a new
layer of precursor is formed next to it. In the course of tooth
formation, differential deposition of dentin and enamel creates the
proper shape of the tooth. This specimen comes from a mouse fetus,
and it's early enough in fetal development that no actual calcification
of the enamel has yet occurred, hence it's really "pre-enamel." Dentin
calcification hasn't yet taken place, so pre-dentin is visible.
The odontoblasts live in a nice protected environment and continue
their activity throughout life. The walls of the pulp cavity are made of
dentin, of course, lined by the active odontoblasts. The inductive
stimulus the ameloblasts need
comes from the CT overlying
them as the tooth bud
forms. This material, the stellate
reticulum, originates as part of
the mesenchyme of the head.
Once the tooth erupts through
the gumline, the cells of the
stellate reticulum die, the
stimulus is lost, hence the
ameloblasts die. Enamel can't
be made anew* and injuries to it
are permanent.

Enamel is hard but brittle. the

retreat of the ameloblasts
leaves small channels through
its substance, which are weak
points. An impact can cause a
fracture along these planes of
cleavage, and if it's bad enough
it can expose the underlying
dentin. Damage to the enamel
can also occur through dietary
exposure and bacterial
infection. If the enamel is
damaged and not repaired, the dentin in turn is subject to mechanical
injury or erosion. Eventually, the dentin will be perforated and the
pulp cavity exposed to infection. This will result in serious problems;
in addition to almost certain loss of the tooth, a really bad infection
can spread to the surrounding jaw bone.

The image at left is from an 18-day mouse fetus, once that was very
close to term (21 days in mice). It's a molar. It hasn't yet erupted, but
it's close to the gum, and had this mouse been born, shortly
afterwards the tooth would have broken through. The enamel is gone
but the space where it was (E) is marked. This is a processing artifact:
teeth have to be decalcified before cutting. Dentin persists because it
has much more organic material than mineralization, so something is
left behind to take the stain.

The complex shape of this tooth indicates how exquisitely the

process is coordinated. The ebb and flow of inductive signals in the
formation of the hard materials actually sculpts the proper shape for
the tooth in question, be it a molar, a canine, or an incisor. The root
formation is similarly controlled by intercellular signaling that
switches on and off as needed; and by selective timed losses of cell
populations and sub-populations. How this process is directed is not
understood except in the most general sense.

*In some animals, especially in rodents, a portion of the enamel

making apparatus remains viable below the gumline, and in these new
enamel is made continuously. For rodents this provides a fresh,
sharp, chisel-like edge to the front of the incisors. That's why they
gnaw continuously: to wear away the excess and keep the edge
sharp. If they are stopped from chewing their teeth will continue to
grow until they puncture the roof of the mouth.

Lab Exercise List